NUYTSIA
Bulletin
of the Western Australian
Herbarium
Vol. 1 No. 3
1972
Department of Agriculture of Western Australia
97093
NUYTSIA
Bulletin
of the Western Australian
Herbarium
Vol. I No. 3
1972
Department of Agriculture of Western Australia
97093 -{ 1 }
A revision of the Australian species of Hybanthus Jacquin
(Violaceae)
By Eleanor M. Bennett
Abstract
A taxonomic treatment is presented of the Australian species of Hybanlluis.
The history of the genus and the morphology and anatomy are discussed. The
Australian species all belong to the subgenus lonidium Schulze, two being in the
Section Snffruticosi Schulze and the other nine in the new Section Variabiles
proposed here. The following new taxa and new combinations are proposed:
H. enneaspermus (L.) F. Mueil. subsp. stellarioides (Domin) stat. nov., H.
florihundiis (Lindl.) F. Muell. subsp. adpressiis subsp. nov., H. floribiindus subsp.
cwvifolius subsp. nov., H. voliibilis sp. nov. and H. vernomi (F. Muell.) F. Muell.
subsp. scaber subsp. nov.
Chromosome counts are given for 8 of the 1 1 Australian species and the
cytology is discussed. Distribution maps for all taxa are given.
Introduction
The name Hyhcmtbus was published by Jacquin in 1 760. It was based on
the single species H. Iiavanensis, which is therefore the type. St. Hilaire (1824,
n.v.), united Hybanthus with lonidium Vent. (1803), as the latter name was in
current use and a renaming under Hybanthus would have caused too much
confusion. In 1876 Mueller transferred the then accepted Australian lonidium
species to Hybanthus,
At the Vienna Congress in 1905 the name Hybanthus Jacq. (1760) was
conserved against its earlier synonym Co/cco/a/va Loefl. (1758). This con-
servation was later considered to be unnecessary as it was thought that the
latter name was invalid being unaccompanied by a description, but Dandy
(1969) pointed out that a generic description was provided. Therefore Cal-
eolaria Loefl. must remain on the list of nomina rejicienda.
According to Airy Shaw (1966) there are 150 species of Hybanthus in
North America, South America. Australia, Malaysia, South Africa and Asia.
Eleven species occur in the mainland states of Australia, none being recorded in
Tasmania.
Morphology of the Australian species
Habit
All the Australian species of Hybanthus develop secondary thickening,
most being shrubs ranging from a height of 50 cm in the bushlets of H. epa-
croides to 2 m in the inland form of H . floribundus. One species, H. epacroides,
develops spinescent branches. The American species H. havanensis also
develops these spines, a feature comparatively rare in the Violaceae.
Hybanthus calycinus, H. debilissimus, H. rolubilis and H. monopetalus are
perennial herbs developing aerial growth from an underground rhizome. With
age the rhizome and base of the stems develop a white pithy bark. This bark
also develops on the stems of H. aurantiacus where it becomes noticeably
fissured with age.
Leaves
The leaves of most species are alternate, but H. monopetalus has opposite
leaves at the upper nodes in addition to the alternate leaves of the lower nodes.
H. bilobus and H. epacroides have leaves clustered at the nodes.
218
Typically the margins are entire, exceptions being the denticulate margins
in H. aurantiacus and crenate margins in some plants of H. enneaspermus
siibsp. stellarioides. The teeth and leaf apices are all tipped with a deciduous
ovoid gland.
A pair of stipules is generally found at the base of each leaf. These may
persist but are usually deciduous. One species, //. dehUissimus, has no stipules.
Inflorescence
This is one of the most variable structures within the Australian species ol
Hyhanthns. Some species have solitary flowers, others have cymes or racemes.
Schulze (1936) believed that the single axillary flowers of Hybanthus
species arose through the gradual reduction of an axillary cymose inflorescence.
This is the view commonly accepted and is illustrated by Lawrence (1951) and
Rickett (1944, 1955).
In the Australian species the inllorcscence types range from single flowers
in the axils of leaves through cymes to racemes. Generally the inflorescence
is axillary but occasionally it may be terminal. //. cynndosus has present on
the same plant dichasial inllorescences as well as various modicalions but the
llowers are never solitary. H. Jlorihundiis shows variation Irom a dichasium to
a solitary flower. In several plants the first division is dichasial, but in many
of the subsequent bracts only one of the pair will develop a flower, the other
bract containing an undeveloped bud. If further growth is monochasial
(racemose) a short cincinnus is produced.
Peduncle and Pedicel
The peduncle of a solitary flower is in this paper considered to be the length
of the flowering stalk occurring below the pair of bracts, and the pedicel is
the portion above the bracts. These vary in relation to each other in length
and pubescence.
The modified leaves on the flowering stalk are therefore correctly called
bracts as they terminate the peduncle. They occur in pairs but one of the pair
is situated slightly higher than the other. The flower buds are in the axils of
the bracts.
Anterior Petal
This petal in the Australian species is distinctly spurred and is much larger
than the lateral petals.
The lamina is typically suborbicular varying from ovate to reniform, and
is generally blue-violet, except in H. aurantiacus and //. enneaspermus subsp.
nellarioides which have yellow flowers. Veining of a darker colouration is
present in the blue-violet flowering species.
The petal is distinctly clawed and the claw may be equal to or less than the
length of the lateral petals. Generally the claw is white with pale blue or yellow
areas. Two parallel raised ridges extend the length of the claw and into the
proximal part of the lamina.
Between the ridges are varying amounts of white pubescence which may
extend into the spur. The presence of the ridges and pubescence probably
helps to prevent the loss of the secretion from the nectaries which are at the
base of the anterior anthers.
Lateral Petals
There are two pairs of differently shaped petals of approximately equal
length. The two outer petals are usually 1 -nerved and narrower than the two
inner petals which may be 3-nerved as in H. vernonii or 5-nerved as in H.
calycinus.
219
Figure 1- Anther structure (diagrammatic). I— Under surface of posterior anther. 2— Upper
surface of anterior anther. T — terminal connective appendage; L = loculi appendages;
C connecting tissue; B = bilocular anther; A = anther surface which may be glabrous or
have varying degrees of pubescence; G basal gland.
The petals of the outer pair are typically narrow-oblong and entire. The
petals of the inner pair show numerous variations in shape between the species,
but typically they are up to 0-5 mm longer than the outer petals.
Androecium
All five stamens are free with the edges of the anthers apparently united by
connecting tissue. All have conspicuous terminal connective appendages
(Figure I) which arc imbricate making the anthers appear to be connate into a
tube. In all species the stamens are shorter than or equal to the lateral petals.
In all species the anterior pair of stamens bear at the base spur append-
ages which are nectaries. These nectaries may be sessile and glabrous as in
H. monopetalus, glabrous and stalked as in some forms of H. etmeaspennus,
stalked with a tuft of hairs at the tip as in other forms of H. enneaspermus, or
densely pubescent as in H. aurantiacus.
Apical loculi appendages (Figure 1) are found in some species or in some
forms within a species. These appendages are narrow-triangular and always
less than 0-5 mm long.
Gynoeciiim
The ovary, composed of three carpels, is regular and unilocular. It is
distinctly 3-angled and in young fruit appears to be 3-keeled. In all species the
ovary is ovoid-globose and glabrous. The style is two to three times as long as
the ovary, beak shaped, and orientated towards the front of the flower.
220
Capsule
The capsule is thick-walled, coriaceous and distinctly 3-sided. It dehisces
by splitting elastically from the apex into the three carpel constituents, the seeds
being shot out. Some capsules of H. calycimts when left in the laboratory
catapulted seeds up to a distance of one metre.
Seeds
The seeds of most species are ovoid-ellipsoidal and dark brown, ranging
in size from 1 -5 mm in H. dehilissinnims to 5 mm in H. calycinus. However,
the seeds of //. enneaspermus and H. aurantiaciis are elongate-ellipsoidal and
yellowish in colour.
When soaked in water the seeds of H. enneaspermus and //. aurantiacus
exude a mucilaginous substance. This was not seen when seeds of the other
species were immersed in water.
The seeds of all species develop a thick, white, fleshy arillus, which, accord-
ing to Raju (1958), originates as a small lump on the raphe as a result of divi-
sions in tire epidermal and hypodermal layers.
Following the definitions given by Martin (1946) the embryos of H.
enneaspermus and H. aurantiacus are regarded as being spathulate whereas the
embryos of the other species are linear. Martin found that in Viola eriocarpa
and Hyhanlhus coneolor the embryos were also spathulate. In H. enneaspermus
and H. aurantiacus the embryo represents approximately half unit volumetric
proportion of embryo to endosperm, whereas in the other species the embryo
represents approximately one quarter unit.
Chromosome numbers (Table 1)
Chromosome counts have been made of 8 of the 10 Australian Hyhanthus
species including 7 of the 10 subspecies recognised. Most counts were obtained
from pollen-mother cells which were fixed in acetic alcohol, stored in absolute
alcohol and stained in acetic orcein. A few counts were obtained from root
and shoot tip material, this previously having been treated with 01 per cent
colchicine and macerated in HC I -alcohol before staining in acetic orcein.
The chromosomes at metaphase I in Hyhanthus monopetalus are much
larger than in any other species, but no mitotic material was available. Gen-
erally the mitotic material examined was unsatisfactory for karyotype analysis,
for the chromosomes are so small that differences in appearance or size cannot
be detected.
The chromosome numbers found are n 4, 6, 8, 12, 24. The base num-
bers of X 4 and x 6 must be accepted for the genus. Four species have the
base number x - 4. Two of these, H. enneaspermus and //. aurantiacus, are
included in the section Suffruticosi and the other two, H. monopetalus and
H. voluhilis, are in the section Variahiles, this separation being based on mor-
phological and anatomical features.
Hvhanihus ftorihundus has a wide range of haploid numbers (n 6, 12, 24).
The plants with n 24 occur as a small population in which only about 6 plants
were found. They cannot be distinguished morphologically from n = 12 plants
collected from other localities. Development and settlement in the area has
probably affected distribution and this polyploidy may well be a result of iso-
lation. No work was carried out to see if the plants in this population are
apomitic or inbreeding.
Although chromosome numbers of n - 6 and n = 12 occur in plants of
H. caivcinus, there appear to be no morphological distinctions between the
cytotypes.
221
TABLE 1
Chromosome number of Hybanthus species.
Species
Locality
n
2n
Collection Number
Subgenus lonidiiim
Section Siiff'niticosi
H. enneaspermus
subsp. enneaspermus
King River
Darwin
Broome
8
8
16
EMS 1731
NB 595
L 6001
H. aurantiacus ....
Durack River
8
EMS 1822
Broome
16
EMS 1925
Walter James Range ....
8
ASG 8887
Section Van'abiles
Group I
H. ftoribundus ....
Parkerville
24
EMB 2207 (x2), 2801
subsp. floribundus
Brookton Highway ..
12
24
EMS 532
W. of Merredin
12
EMB214I (x3)
S. of Salmon Gums . ..
12
EMB2156(x3)
*Ravensthorpe
12
24
EMB 2190
S. of Pingrup
12
EMB 2199 (x2)
Kalgarin
12
EMB 2201
E. of Kondinin
12
EMB 2202 (x2)
W. of Kondinin
12
EMB 2203
E. of Corrigin
12
EMB 2204
Geraldton
12
EMB 2241 (x5)
Perenjori-Paynes Find
12
EMB 2243 (x4)
Warriedar Station
12
EMB 2245
Dongara
12
EMB 2216
Kalbarri
12
PGW 6725
E. of Southern Cross
6
12
EMB 2151, 2803
H. ftoribundus ....
Norseman
6
12
EMB 2152 (x3)
subsp. curvifolius
Baliadonia
6
PGW 7738
S. of Coolgardie
12
EMB 2804 (x2)
H. ftoribundus ....
E. of Ravensthorpe ...
6
EMB 2181 (.x2)
subsp. adpressus
Ravensthorpe
6
12
EMB 2187 (x5), 2455,
2541
Ravensthorpe
12
RAS 512
Hopetoun Road
6
EMB 2195 (x2)
S. of Ravensthorpe ...
6
EMB 2189
H. cymulosus
Mt. Singleton
6
EMB 2246-2249
Group n
H. monopetaius ....
Culoul Range
4
NSW 99494
Ocean Beach
4
NSW 112261
H. caiycinus
Wyadup
12
EMS 1640
Busselton
12
EMB 2266
Cape Naturaliste
12
24
EMB 2269
S. of Mandurah
12
24
EMB 2254-2257
Naval Base
12
ASG 9190
Manning
12
EMB 2253 1x2)
South Perth
12
24
OAB
Yallingup
12
EMS 1642, 1645
Gnangara
12
EMS 538
Yanchep
12
EMS 510, 551, 523, 525
Lancelin
12
EMS 1332
Moore River
12
TEHA 3068
Hill River
12
TEH A 3069
Dongara
12
EMB 2214 (x2), 2215
(x2)
Dongara
12
24
TEHA s.n.
N. of Northampton ....
6
ASG 9182
NW Coastal Highway
6
EMB 2233 (x4)
N. of Northampton ....
6
EMB 2236 (x4)
222
Species
Locality
n
2n Collection
Group III
H. volubilis
Osmington
8 EMB 2815
H. epacroides ....
W. of Merredin
12
EMB 2139 (x3), 2140
S. of Salmon Gums ..
12
EMB 2166
H. hilobus
N. of Esperance
12
EMB 2167 (x3)
Salmon Gums
12
EMB 2168
E. of Esperance
24
PGW 8076
Ext! a- A usual fan Niimhers
Species
Number
Country
Reseacher
Source of
n 2n
Reference
Subgenus tunidium
Section Siiffriilicosi
H. parviflonis
12 24
South .'Xmerica
Heilborn
Darlington and
Wyle (1955)
H. enneaspermus ....
32
Tropical Africa
Mangenot, S. &
G. Mangenot
OrndufT (1967)
* Large white-flowered plant growing among n 6 plants of subsp. adpressus.
The count of 2n -32 for the tropical collection of Hyhanthus enneaspermus
indicates that it is a tetraploid if compared with the Australian forms of the
species. A count of 2n 24 has been obtained for the South American species
Hybantinis parviflonis (Darlington and Wylie 1955). This agrees with the base
number x 6 obtained for some ot the Australian species.
Pollen
The tetrads of Hyhanlhus calyciiws, H. florihundus, H. cymulosus, H.
epacroides, H. enneaspermus subsp. enneaspermus and H. monopetalus are
decussate or tetrahedral. All cells have four developing pollen grains, but many
also contain one or two microcytes.
The pollen fertility shows no correlation with chromosome number.
Hyhantbus caivcinus and H. florihundus, irrespective of whether n 6, 12 or 24,
record fertility of up to 95 per cent, whereas other species record a high sterility.
In H. ealvcinus and H. florihundus the pollen of n 12 plants is significantly
larger than the pollen of n 6 plants, but the n=24 plants have pollen well
within the size range of the n 12 plants.
Anatomy
The material for anatomical work was fixed in FAA and sectioned with a
hand-operated bench microtome. Several samples were also embedded in wax
before cutting sections with a rotary microtome. The prepared sections were
stained in safranin in 50 per cent alcohol, and in fast green in clove oil. Leaves
for examination were cleared using the lactic acid technique.
Crystals
Calcium oxalate crystals occur in the leaves of all the Australian species.
223
Borodin in Solereder (1908) distinguished eight different cases (type and
position) of crystals in the Violaceae. Examples of Viola type and lonidium
type have been found in the species examined. The northern species Hyhmthus
auranliacus and H. enneaspermus have the lonidium crystal type, consisting of
clinorrhombic crystals scattered along the veins of the leaves.
All the other Australian species have the Viola-type of crystals in which the
scattered cells contained clustered crystals of druses. These crystals are
scattered in the mesophyll of leaves and in the parenchyma of stem, root and
rhizome.
Hairs
Unicellular hairs occur on some plants of most species, mainly as epi-
dermal cells of the leaf and young stem. Multicellular hairs occur on the stem
of H. floribundus subsp. adpressus. The unicellular hairs vary among species
mainly in the relation of hair length to cell size.
Leaf Anatomy
The leaves of all species are dorsiventral and have a thin cuticle. The
epidermis is one-layered except in H. enneaspermus where there are layers over
the midvein. The epidermal cells of this species contain mucilage. In all
species stomata, mainly of the cruciferous type, occur on both the upper and
lower epidermis.
In H. enneaspermus and H. aurantiacus the rectangular crystals are dis-
tributed along the veins. In all the other species the crystals are druses scattered
throughout the mesophyll.
Stem
In all the species examined the stem structure is that of a typical dicotyle-
don. With age the epidermis of the stems develops a bark layer of several
rows of periderm cells.
In all the Western Australian species except H. volubilis the parenchyma
cells of the cortex have scattered included crystals. Phloem fibres are developed
in all species. These are not conspicuous in young stems but in the older
stems they develop as a band several layers thick. H. volubilis in cross section
appears to have a triangular stem with a bundle of fibres well developed below
the angles which continue as a 1-or 2-celled layer around the remainder of the
stele.
In all species the central pith layer consists of parenchyma cells; some of
these have included crystals which do not, however, occur in H. volubilis.
The vessels are spirally thickened and have opposite bordered pits. In H.
epacroides and H. bilobus the pits are simple. The perforation plates are
simple and the intervascular pitting is opposite. The fibres are septate with
simple pits in H. epacroides and H. bilobus, and with bordered pits in the other
species.
Nectaries
Only the sessile nectaries in the Hybanthus species from the South-West
Land Divisions were examined. Scattered stomata occur on the nectary sur-
face, particularly towards the base. The outer cells of the nectary are smaller
than the remaining secretory cells. Covering the cells is a cuticle. The
remainder of the gland consists of numerous secretory cells closely packed
together in a dense mass.
Citation of Specimens
In the citation, the districts in Western Australia are those given by Diels
(1904). For most species only a few of the herbarium specimens seen have been
cited. The abbreviations used for herbaria follow the Index Herbariorium pt.
224
1 ed. 5 (1964), with the addition of GAUBA for the Department of Botany,
the School of General Studies, Australian National University, Canberra, and
UWA for the Department of Botany, University of Western Australia.
HYBANTHUS
Hyhanthiis Jacq., Enum. Plant. Carib. 2:17 (1760), nom. cons. Type: H. havanensis Jacq.
Calceolaria Loefl., Iter Hisp. 183 (1758), nom. rej. Type: C. calceolatia Loefl.
? Pomhalia Vand., Fasc. PI., 7 (1771). Type: P. ipecacuanha (L.) Vand. ex T.
Soica Spreng. in Schrader, Joiirn. Fur die Bot, 2:192 (1802). Type: S. vcriicillata (Orteg.)
Spreng.
lonidium Vent., Jard. Malmai.son t, 27 (1803). (nom. illeg., nomenclatural synonym of Soiea
Spreng.). Type: /. pnlyi’ulaefoliiim Vent. /. verlicillafa (Orteg.) Roem. et .Schult.
y Cuheiium Raf., Catalog. 13 (1824). Type: C. concoior (T. F. Forster) Raf.
Pigea Ging. in DC., Prodr. 1 :307 (1824). Type: P. filifurniis Ging. in DC.
Ionia Pers. ex Steud., Nom. Bot. ed I. 1 :433 (1821) lonidium
Vlamingia De Vr. in Lehmann, PI. Preiss. 1:398 (1845). Type: F. australasiaca De Vr.
y Acentra Phil.. Sert. Mendoc. alt. 3 (1871). Type: A serrala Phil.
Clclamlia J. M. Black, Journ. and Proc. Roy. Soc. S. Austral. 56:46 (1932). Type: C. con-
vaUis J. M. Black
y Orthion Standi, et Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 22:249 (1940). Type:
O suhses.sile (Standi.) Standi, et Steyerm.
Not all the above generic names have been applied to Australian Hybanthus
species. However, Australian species have been described or included in
Calceolaria, Soiea, fonuliiari, Pigea, Vlamingia and Clelaiulia. These six
genera I consider to be synonyms of Hyhanllnis, an opinion arrived at from the
study of the type species or of their descriptions.
It is impossible for me to draw conclusions about the synonymy of the
other genera, particularly as there is disagreement in recent literature. Mel-
chior (1925) included the first eleven genera under Ilyhanllnis. Hutchinson
(1967) considered Cuheiium to be a distinct genus, but all the other genera he
regarded as synonyms of Hybanthus. Airy Shaw (1966) regarded Clelaiulia
and Orthion as separate genera, but all the other genera, including Cuheiium
he regarded as synonyms of Hybanthus.
Perennial herbs, undershrubs or semi-creepers. White pithy bark dev-
eloped on underground rhizome of perennial herbs and on tap roots of under-
shrubs. Stems branching at base or above, smooth or ribbed from leaf bases
and stipules, glabrous to densely pubescent, developing thin or pithy bark with
age. Leaves alternate or clustered, occasionally opposite, sessile or shortly
petiolate, linear to lanceolate, glabrous or with scattered to dense pubescence of
simple hairs, sometimes tomentum disappearing with age, flat to conspicuously
revolute; margins entire or crenate or dentate; tip acute or obtuse. Stipules
present or absent, terete or flat and linear to lanceolate, entire or divided, white
or grey or green, all tipped with prominent usually deciduous gland. In-
florescence axillary, cymose or racemose, or flowers solitary. Peduncle and
pedicel approximately equal in length; bracts, coriaceous or hyaline, glabrous
to densely pubescent or ciliate along margins, green, or grey or white. Flowers
zygomorphic. Sepals 5, free, subequal, linear to ovate, blue to green, margins
entire or ciliate. apex recurved or appressed to petals. Corolla of 5 free petals,
violet to purple, or yellow to orange. Anterior petal distinctly spurred and
prominently clawed between lamina and spur; claw with two parallel ridges
which extend into both the spur and into the prominently veined lamina. Lateral
petals of dissimilar pairs; 2 outer petals linear, prominently 1-3 nerved: 2
inner petals broader, lanceolate, ovate or falcate-oblique, prominently nerved.
Stamens 5: filaments short; anthers free (but appearing united by apical and
lateral connecting tissues), narrow ovate, erect, bilocular, introrse. dehiscing in
longitudinal slits, glabrous or penicillate with long white hairs, or with scattered
to dense short white pubescence, sometimes bearing orange or white subulate
loculi-appendages; nectaries at base of two anterior anthers.
225
Ovary superior, ovoid-globose, tricarpellate, unilocular, placentation parietal,
ovules 3-15; style simple, sigmoid; stigma expanded above, orientated towards
front of flower. Fruit a thick-walled coriaceous capsule, splitting in three at
dehiscence, subtended by persistent perianth. Seeds 1-12, globose to oblong,
surface reticulate, foveate, ribbed or smooth. Embryo spathulate or linear,
endosperm copious, dark brown or yellow.
TABLE 2
Comparative features of the Sections Suffruticosi and Variabiles.
Section
Suffruticosi
Variabiles
Distribution
Northern Australia
Southern Australia
Flower colour
blue, yellow, orange
mauve to white
Seed colour
yellowish white
dark brown
Testa surface
ribbed, foveate or smooth
smooth, foveolate or reti-
culate
Nectaries
H. eiineaspernuis, elongated,
glabrous or penicillate; H.
aurantiacus sessile, hirsute
sessile, glabrous
Ovules
12^
6 or less
Embryo
spathulate
linear
Chromosome number
n = 8
n - 4, 6, 12. 24
Crystal shape distribution ....
rectangular, along veins
druses, scattered in leaf,
mesophyll, root and
stem parenchyma
Venation
close reticulate ....
open reticulate
Infrageneric di visions
Schulze (1936) divided the American species of Hvbanthus into the two
subgenera Euhybanlhus and lonidiuny the former having’the filaments and to a
lesser extent the anthers connate into a tube whilst in the latter they are free.
All the Australian species have free anthers and so are included in the sub-
genus lonidium.
This subgenus he divided into four sections, three of which are not appli-
cable to the Australian species but the section Suffruticosi includes the two
northern Australian species H. enneaspermus and H. aurantiacus. Schulze
recognised that a new section was required to provide for the remaining eight
Australian species. This section I propose to call Variabiles due to the variation
m the inflorescence forms found within the group. A comparison of the sec-
tions Suffruticosi and Variabiles is given in Table 2.
The Variabiles are characterised by the following features
(i) shrubs or perennial herbs.
(ii) leaves alternate, although one species has opposite leaves in the
upper part of the plant.
(iii) base of anterior petal distinctly gibbous.
(iv) nectaries at the base of the anterior stamens flat and sessile
(v) ovules 6 or le.ss in the ovary.
The section can be further divided into three groups on the form of the
inflorescence.
226
Group 1. Inflorescence dichasial: H. fiorihiiiuliis and H. cymiilosus.
Group 11. Inflorescence a raceme; H. monopetaliis, H. dehilissimus and
H. calycinus.
Group 111. Flowers solitary: H. vernonii, H. voluhilis, H. epacroides and
H. hilohus.
2. H. aurantiacus
8 .
9.
H. voluhilis
H. vernonii
Key to the Australian Hybanthus species
1. Flowers solitary, subtended by one pair of bracts.
2. Inner lateral petals lanceolate falcate.
3. Gland at base of anterior anthers elongated, glabrous or tipped with tuft of long hairs
1. H. enneaspcrmus
3. Gland at base of anterior anthers sessile, usually densely hirsute
2. Inner lateral petals oblong.
4. Anterior petal 6 mm long or longer.
5. Plant twining
5. Plant a shrublet. never twining
4. Anterior petal 3-5 mm long.
6. Plant glabrous or with few scattered hairs
6. Plant densely hirsute.
7. Leaves linear; sepals glabrous or sparsely pubescent
7 I eaves obovate to bilobed; sepals densely hirsute along margins and veins
11. H. hilohus
I. Flowers not solitary (or if solitary several pairs of sterile bracts present below flower).
8, Inflorescence racemose.
9. Inner lateral petals lanceolate-falcate
9. Inner lateral petals oblong.
10. Stipules triangular, prominent. Racemes about 20 cm long
10. Stipules absent or very small. Racemes 2-3 cm long
8. Inflorescence dichasial or branched.
1 1 . Stipules flat. Anterior petal 9 mm or more long
II. Stipules terete. Anterior petal less than 9 mm long
4. H. Horihundus
10. H. epacroides
5. H. monopetalus
7. H. calycinus
6. H. dehilissimus
3. H. cymulosus
4. H. Horihundus
Subgenus ionidium G. K. Schulze, Bot Jb 67: 453 (1936). hmidium VenU
.lard. Malmaison t. 27 (1803). Type; /. polygalaefolium Vent. (nom. illeg.) =
/. verticillata (Orfeg.) Roem. el Schult.
Section Suffruticosi G. K. Schulze, Bot. Jb. 67: 453 (1936). Schulze des-
cribed only South American species and as I have not seen any of these species
I feel incompetent to choose a lectotype for the section.
1. Hybanthus enneaspcrmus (L.) F. Muell., Fragm. 10:81 (1876).
Viola enneasperma L.. Sp. PI. 2 : 937 (1753). Type from Ceylon, Hermann (holo. : BM,
photo seen).
For synonymy see Tennant 1962. From a study of the photographs of the
types of Viola suffrutkosa L. and Viola enneasperma L., and because of the
variability noticeable within the species, 1 have decided to follow Tennant and
to regard them as being conspecific.
Perennial spindly herb or compact shrub to 60 cm tall. Stems glabrous
or with ^ scattered hairs. Leaves alternate, subsessile; lamina (5x1) 40x2
(90x5) mm linear to lanceolate, margins entire to crenate, glabrous or with
scattered pubescence; stipules linear to subulate, 1-7 mm long, glabrous, grey-
white Flowers solitary. Sepals lanceolate, 3-4 mm long, acute, + prom-
inently carinate, glabrous or with ; scattered pubescence (especially along
keel) green. Anterior petal 8-15 mm long; lamina violet or pale yellow.
Lateral petals of dissimilar pairs; 2 outer linear-oblong, 3-4 mm long; 2 inner
lanceolate-falcate, 4 -5-5 -5 mm long. Stamens: filaments dimorphous, 3
oosterior short, 2 anterior equal in length to anthers and bearing elongated
glabrous nectaries tipped with a dense tuft of hairs; anthers oblong-elhptic,
loculi appendages absent. Capsule 4-9 mm long; seeds 5-12, ovoid-elhpsoidal,
2-3 mm long, longitudinally ribbed, ± pitted between ribs, yellowish-white.
227
Key to subspecies
I. Stipules long (up to 4 mm). Margins of leaf closely revolute, usually glabrous, but if
pubescent then hairs spreading. Flower blue subsp. enneaspermus
1. Stipules short ( ± I mm). Margins recurved, leaves up to 5 mm wide, hairs always antrorse.
Flowers yellow subsp. stellarioides
subsp. enneaspermus
Pigea banksiana Ging. in DC., Prodr. 1 :307 (1824). Type: “ in Nova-Cambria austr. Viola
angustifolia Banks herb. (DC. vs. in h. Banks).” (holo; BM, photo seen).
loiiiditim banksianum (Ging. in DC.) Steud., Nomen. Bot. Ed. 2. 1:813 (1840).
Hyhanthus enneaspermus var. communis f. angustifolia Domin, Biblioth. Bot. 89:982 (1928).
Type: Lower Victoria River, F. Mueller, May 1856 (lecto: K, photo seen).
Hybanthus enneaspermus var. banksianus (Ging.) Domin, l.c.
Flowers blue. Leaves entire or crenate, margins closely revolute occas-
ionally fiat; I pubescent with spreading hairs, dark green on upper suface,
paler underneath. Stipules up to 4 mm long, scarious, opaque, midvein d
prominent.
Distribiifion: This subspecies is found in the north of Western Australia, the
Northern Territory, Queensland and Northern New South Wales (Figure 2).
In addition it is found in Malaysia, South and East Africa, India and Ceylon.
272 collections were seen from Australia and South Africa.
IVestern Australia: Gibb River, C. A. Gardner 9991 (PERTH); Kimberley District, 1888,
E. Giles (MEL); King River, October 1906, fF. V. Fitzgerald (NSW); Karunjie Station, 6
Feb. 1959, J. B. Ritson (CANB).
Figure 2 -Distribution of Hybanthus enneaspermus (L.) F. Muell. subsp.
228
enneaspermus.
Northern Territory: South Bay, Bickerton Island, R. L. Specht 519 (AD); 64-7 miles SE Top
Spring Store, G. Chippendale 5815 (PERTH); Stoward Springs, 5 Apr. 1958, W. McKay
(BRI); Port Darwin 1883, Foeisch (MEL); 14 miles SE Mountain Valley, D. J. Nelson 234
(NT).
Queensland: 2 miles E of Normanton Township, 5 Mar. 1954, M. Lazarides (NT); Cooktown,
1877, A. W. Persietz (MEL); between Cloncurry and Camooweal, June-December 1899,
R. C. Barton (BRI); Townsville, August 1901, E. Betche (NSW); Bloomfield, IV. L. Jones
1564 (CANB); vicinity of Nicholson River, 1886, Dietrich (AD); Alva Beach, Ayr, R. Carolin
4620 (SYD).
New South IVales: New England near Armidale, Bishop Twiner (MEL),
subsp. stellarioides (Domin) E. M. Bennetl slat. nov.
H. enneaspennus var. stellarioides Domin, Biblioth. Bot. 89:983 (1928). Type: In rupibus
collis apud. opp. Yarraba, alt, cca 550 m s.m. K. Domin 6794, January 1910 (holo: PR).
H. enneaspermus var. communis f. flavus Domin, l .c. Type: " Keppel Bay, R. Brown Iter
Australiense 1802-05 No. 4949 als Pombalia flava.” (holo: K, photo seen).
H. enneaspermus var. communis f. puhescens Domin, l.c. Type: “ Northumberland Islands,
R. Brown No. 4949 als Pombalia flava f. mollis." (holo: n.v.).
Flowers yellow. Leaves entire or crenate, 1 pubescent on margins, hairs
antrorse, dark green on upper surface, paler beneath. Stipules short, adpressed
against stem, short, scarious, opaque, margins pubescent.
Distribution: This subspecies is found close to the east coast of Queensland and
New South Wales (Figure 3). 132 collections were seen.
Queensland: Ml. Cotton, S. L. Everist 1019 (BRI); Broadsound and Heads of the Isaacs.
R. Brown 155 (MEL); Bramnio Bay, Dunk Island, July 1917, E. J. Banfield (NSW); "The
Rocks” Ayr, 13 Mar. 1952, F. H. Kleinsmidt (CANB).
New South Wales: Moona River, Walcha, Dec. 1884, A. R. Crawford (MEL)\ Hathead Mt.
Korogora Pt., 19 Jan. 1953, E. F. Constable (NSW); K.ew, Autumn 1917, J. B. Cleland (AD);
Kendall, Feb. 1950, T. M. White (SYD).
Figure 3— Distribution of Hybanthus enneaspermus (L.) F. Muell. subsp. stellarioides (Domin)
stat. nov.
229
2. Hybanthus aurantiacus (F. Muell. ex Benth.) F. Muell., PI. North West.
Austr. : 5 (1881)
lonidiiim lunantiacum F. Muell. ex Benth., FI. Austr. 1:102 (1863). Type: N.W. Coast of
.Australia, Byiwe (syn; K, photo seen); Victoria River (syn: K, photo seen; MEL); Cygnet
Bay. N. W. Coast A. Cunningham (syn; BM, photo seen).
Hybanthus enneaspennus var. aurantiacus (F. Muell, ex Benth.) F. Muell., PI. Indig Shark
Bay 6 (1883).
Hybanthus miniatus F. Muell. et Tate, Trans. Proc. Roy. Soc. S.A. I3;97 (1890) (nom, nud.).
Type: Gills Creek, S.A., (n.v.). See Trans. Proc. Roy. Soc. S. Austral. 19;83 (1894).
Hybanthus elegans Domin, Biblioth. Bot. 89:984 (1928). Type: between Ashburton and Yule
Rivers. W.A., f. Clement (holo; K, photo seen.).
Compact shnth to 60 cm tall. Stems scabrous or L glabrous. Leaves
alternate or clustered at the nodes, subsessile (4x1) 35x4 (65x5) mm, linear to
lanceolate, coriaceous, margin serrate-dentate occasionally entire, densely hir-
sute or occasionally glabrous; stipules linear to subulate, 1-30 mm long, pubes-
cent or occasionally glabrous, green and leaf-like or yellow-white and pithy.
Flowers solitary. Sepals lanceolate, 3-6 mm long, acute + prominently
carinate or 3-nerved. ciliate, otherwise glabrous, green. Anterior petal 7-15 mm
long; lamina yellow to orange. Lateral petals of dissimilar pairs; 2 outer,
linear-oblong, 3-5 mm long; 2 inner, lanceolate-falcate, 3 -5-6 -5 mm long!
Stamens: filaments dimorphous; 3 posterior short, 2 anterior t equal in length
to anthers and bearing sessile glabrous to densely pubescent nectaries; anthers
oblong-elliptic, loculi appendages absent. Capsule 5-9 mm long; seeds 5-10,
ovoid-ellipsoidal, 2-3 mm long, smooth, pitted, or longitudinally ribbed’
yellowish white.
Distribution: This species is found in the north of Western Australia, Nor-
thern Territory, Queensland, and far north of South Australia (Figure 4).
221 collections were seen.
H'esleni Australia: Cape Range, A. S. George 2508 (PERTH); Forlescue River, June 1878,
./. Forrest (MEL.); Lennard River, May 1906, IT'. K. Fitzgerald (FiSV\/)\ Corong Creek, S. of
Port Hedland, N. T. Burhidge 5838 (CAMB); Pass of the Abencerrages, P. G. Wilson 2437
(AD).
Northern Territory: 50 miles N. ot Tennant Creek township, R. A. Perry 608 (BRI); Alice
Springs, J. R. Maconochie 471 (PERTH); Ormiston Gorge, G. M. Chippendale 2116 (NT);
Mt. Liebig, 23 Jul. 1966,7. H. Willis (MEL); I mile N.W. Lassetters Cave, Peterman Range,
II Mar. 1959, D. J. Nelson (NSW); 9-5 miles NNW Alice Springs township, 5 May 1955.
M. Lazarides (CANB); New Rabunlja, 1932-33, Strehlow (AD); Alice Springs, Gaiiha (CBG);
Emily Gap, 7 Jul. 1966, R. C. Carolin (SYD).
Queensland: Lake Corella, Mary Kathleen, W. ./. Lavery 110 (BRI); Georgina River, Sept.
1910, E. W. Bick (BRI); Clondurry, Flinders River, 1876, A. Henry (MEL); Currawilla,
S. L. Everlsl 3941 (CANB), Moongerrie, 16 Jan. 1928, R. D, Riddell (AD).
South Australia: Cooper Creek, E.N.S. Jackson 428 (AD).
Section Variables E. M. Bennett sect. nov.
Elores lilacini vel albi. Glandulae in antheris anticis sessiles, glabrae. Ovarium ovulis
usque ad 6; embryo linearis.
Type: Hybanthus calycinus (DC. ex Ging.) F. Muell.
Flowers mauve to white. Nectaries on anterior anthers sessile, glabrous.
Ovules 6 or less per ovary; embryo linear. Seeds dark-brown, surface smooth,
foveate, or reticulate. Calcium oxalate druses scattered in leaf mesophyll, and
in root and stem parenchyma.
Group 1 : Inflorescence dichasial.
3. Hybanthus cymulosus C. A. Gardn., Journ. Roy. Soc. W. Austral. 22: 125
(1936).
Type: Austin district, at the Northern base of Mt. Singleton. W.A., W. E. Blackall and C. A.
Gardner, 9 July 1931 (holo; PERTH).
Perennial herb to 70 cm tall. Leaves alternate, sessile, narrow-lanceolate
to linear, 20-50 x 1-3 mm; stipules 0- 1-0-5 mm long, tripartite to laciniate,
occasionally entire. Inflorescence 3-to many-flowered in axillary dichasia, 2-5
cm long. Sepals lanceolate 4-6 mm long, tip slightly recurved, ciliate on
margin, mid nerve prominent, blue to green. Anterior petal 8-15 mm long,
sparsely ciliate on edge, pale violet. Lateral petals of dissimilar pairs, 2-4 mm
long, pale violet ; 2 outer oblong, 1 -nerved, 2 inner orbicular, 3-nerved. Stamens
shorter than lateral petals, no loculi appendages present. Capsule 5 mm long,
seeds 1-3, ovoid-ellipsoidal, ca 3 mm long, dark-brown, prominently pusticulate.
Distribution: Austin District of Western Australia near the townsite of Paynes
Find (Figure 8). 8 collections were seen.
Western Australia: Ninghan, between homestead and Great Northern Highway, A. S. George
3713 (PERTH); 220 miles Paynes Find Road, C. A. Gardner 10209 (PERTH); Northern
slopes of Mt. Singleton, E. A/. Scn-m^-eo/u- 21 12 (PERTH); 7 miles from Great Northern High-
way to Mt. Singleton, E. M. Bennett 2246, 2247 (PERTH).
4. Hybanthus floribundus (Lindl.) F. Muell., Fragm. 10: 81 (1876).
Pigea florihunda Lindl. in Mitch., Three Exped, 2:164 (1838) n.v., ed. 2, 2:165 (1839). Type:
Mitchell’s Journey. 1836, July 6 No. 223 (iso: MEL).
lonidium floribundum (Lindl.) Walp., Repert. Bot. Syst. 2:767 (1843).
Ca/ceo/an'a/7ur/6««r/a (Lindl.) O. Kuntze. Rev. Gen. PI. 1:41 (189U.
lonidium australasiae Behr, Linnaea 20:629 (1847). Type: Sud A.ustrahen Lyndock Valley
in der Barossa range und in dem Pine Forest-Walde, H. Behr. (holo. HAL).
231
lomdium multiflorum Turcz., Bull. Soc. Nat. Mosc. 27, Sect. 2:340 (1854). Type: Nova
Hollandia. Drummond 5X\\ Coll. No. 72 (holo: n.v.)
lomdium hrevHahre Benth.. FI. Austr. 1:102 (1863). Type: West. Austr. Drummond 665
(syn: MEL); Swan River, Drummond, 1st Coll, (syn: n.v.).
Clelandia convallis .1. M. Black, Trans, and Proc. Roy. Soc. S. Austral. 56:47 (1932). Type:
Wilpena Pound, S.A., T. B. Clelaiid (ho\o : AD).
SImih to 1 - 5 m tall. Leaves alternate, sessile, linear, lanceolate or oblong,
5-35 X 0-5-7 mm, acute or obtuse ± recurved; stipules linear 0-5-1 -5 mm long,
glabrous or with a few' trichomes towards base. Inflorescence axillary, dichasial
or racemose to 40 mm. occasionally flowers solitary. Sepals 0 - 75 to 4 mm long,
pale blue to white or dark blue to green; 2 inner petaloid, ± I rnm longer than
3 outer. Anterior petal 4-1 1 mm long, pale blue to white. Lateral petals of
dissimilar pairs: 2 outer petals oblong, 1 -5-3 mm long, l-nerved, 2 inner petals
broadly oblong. 2-5-4 mm long, 3 or more nerved. Anther surfaces glabrous
or with scattered white hairs, loculi appendages sometimes present. Capsule
to 8 mm long; seeds 1-4, ovoid-ellipsoidal, 2-3 mm long, reticulate, foveate, or
smooth, dark brown.
Key to subspecies
I . Sepals petaloid, pale blue, apices recurved. Leaves _ appressed upwards against stem.
2. Leaves narrow, conduplicate (Ravensthorpe area) subsp. adpressus
2. Leaves broad, flat, distinct pustules on surface subsp. floribundus
I. Sepals dark blue to green, apices appressed to petals. Leaves variously spreading.
3. Leaves conduplicate, curved (Norseman area) subsp. curvifolius
3. Leaves flat, not curved subsp. floribundus
subsp. floribundus
Leaves i flat, apex uncinate.
Distribution: This subspecies is found in the south of Western Australia, South
Australia, Victoria and southern New South Wales (Figure 5). 319 collections
were seen.
Western Australia: 12 km SW Queen Victoria Spring, D. W. GWa// 2964 (PERTH); Victoria
Desert Camp, R. Helms 58 (MEL); Cundeelee Mission, March 1963, M. C. George (NSW)-
East Yuna, A C Burns 97 (PERTH); ca. 20 miles east of Merredin, R. Carolin 3089 (SYD.);
Yilgarn, Herb. F Mueller (AD); Wedgicarrup W. of Wagin, N. T. Burhidge 2359 (CANB)-
(CANB); W.A., Sept 1930, E. Ashbv (ADW); Waroonai
July 1907. G. F. Sa/-//iour/( NSW). k , . ^ uuna.
South Unco\n, 1874. /. L. Browne 5 (MEL); Eyre Peninsula, 28 Aug. 1958,
Gauha (OAUBA); Old Coach Road 3-4 miles W. of Stony Pinch. D. E. Svmon 3819 (CANB)-
Wilpena Pound July 1937, .4 F. V. Richardson (ADW); near Chain of Ponds, Torrens Gorge
24 Aug. 1946, y. K(cA-er>- (NSW); Mt. Remarkable, 17 Sept. 1958. H. M. Cooper (ROY, Mt
Lolty Range, y. G. F. j'y/we (BRI).
6 Jul. 1966, C. W. E. Moore (CANB); Stawell, Apr.
I ’ 'Ik (NSW); 12 miles S. of Ouyen, 8 Sept. 1961, M. E. Phillips (CBG)- Warrackna-
beal, 10 May 1903, F. M. Reader (MEL); Bendigo, August (BRI)
'miff'’ 5c«"»"r/(SYD. CANB); near Ardlethan,
8 Aug. 1966, L. D. Pryor (GAUBA); Nericon near Griffith, G. R. Saintiv 257 (NSW)
0«mt.vW.- Queensland. ATarte)- (BRI). This locality Is doubtful.
Included under subsp. floribundus are plants which at their extremes appear
to be subspecifically distinct but which are in fact connected by intermediate
variants.
There is a large form at Queen Victoria Spring in Western Australia which
reaches a much greater height (up to 1 - 5 m) than the common shrubby plants of
this species Two characters were found useful in recognising herbarium
specimens of this plant, firstly the flower size and secondly the glaucous and
pustular appearance of the leaf surface. However plants occurring in South
Australia, particularly those growing around Wilpena Pound, have flowers
which are within the lower size limits of the Queen Victoria Spring plants.
Also some South Australian plants have a leaf texture and surface shnilar to
V^aoHa''*^ collected in Western Australia, South Australia and
232
Plants with oval glaucous leaves occur in various localities throughout
Australia but there are several collections showing intermediates between the
typical and glaucous ones. One of the glaucous plants collected near Southern
Cross has a different chromosome number (n - 6) from the typical subsp.
florihimdus (n 12).
From herbarium specimens alone it is difficult to distinguish these several
different forms; further collecting and possibly cytological work will need to be
carried out before any definite conclusion on their taxonomic status can be
reached.
subsp. adpressus E. M. Bennett subsp. nov.
Folia ad caulem sursum L adpressa, angusta, conduplicata. Sepala petaloidea apicibus
recurvis.
Leaves appressed upwards against stem, narrow, conduplicate. Sepals
petaloid, apices recurved.
Type: 5 miles south of Ravensthorpe, in heavy brown clay, W.A., 16 May, 1968. £. M.
Bennett 2195 (holo: PERTH).
Distribution: This subspecies is found in the Eyre district of Western Australia
near Ravensthorpe (Figure 5). 29 collections were seen.
Western Australia: South-west coast and interior of Western Australia, G. Maxwell (MEL);
I -5 miles along old road north of Ravensthorpe, F. iMlIfitz 5252 (PERTH); 34 miles E. of
Ravensthorpe E. M. Bennett 2189 (PERTH).
Figure 5 — Distribution of Hvbanthus florihimdus (Lindl.) F. Muell. subsp. flcribimdiis (®),
subsp. adpressus E. M. Bennet (o) and subsp. ciirvifoHus E. M. Bennett (x).
233
subsp. curvifolius E. M. Bennett subsp. nov.
Folia conduplicata, manifeste curvata, apicibus uncinatis.
Type: South edge of Lake Cowan, Norseman district, W.A., 15 May 1968. E. M. Bennett
2152. (holo: PERTH).
Leaves conduplicate, distinctly curved, apex uncinate.
Distribution: This subspecies is found in the Coolgardie district of Western
Australia between and east of the townsites of Kalgoorlie and Norseman
(Figure 5). 43 collections were seen.
Western Australia: Comet Vale, Aug. 1917, ./. L. Jutson (MEL); Coolgardie, July 1899, R.
Helms (NSW); Ml. Thirsty, Norseman, J. Barnes 6079 (PERTH); Between the bank and
Eyres Relief, Maxwell (MEL); 13 miles SE of Coolgardie on Coolgardie Esperance Highway,
B. G. Briggs 278 (NSW); Londonderry, K. Newbey 2588 (PERTH).
Group II: Inflorescence a raceme
5. Hybanthus monopetalus (Roem. et Schult.) Domin, Biblioth. Botanica
89: 984 (1928).
loniditim monopetahnn Roem. et Schult., Syst. Veg. 5:400 (1819). Type: “ Ex anglia habuit
Rbmer in Herbario suo.” (holo: probabI> once at LZ, if so now destroyed: ?iso: BM).
Pigea monopetala {Koftm. et Schult.) DC. ex Ging. in DC., Prodr. 1 :307 (1824).
Solea monopetala (Roem. et. Schult.) Spreng., Syst. Veg. 1:804 (1825).
Calceolaria monopetala (Roem. et Schult.) Britten in Banks and Solander. Illustrations of the
Botany of Captain Cook’s Voyage 1 :7 (1900).
Hybanthus monopetalus rar. normalis Domin, Biblioth. Bot. 89:985 (1928) nom. illeg.
Hybanthus monopetalus var ahbreviatus Domin, l.c. Type: Sandsteinhugel der Blue Mts.
nichl selten (Domin IV. 1910); Blue Mts., C. T. White: Port Jackson (Georges Head), R.
Brown 1805; R. Brown Iter Australiense 1802-05 No. 4951; Blue Mts., A. Cunningham, (syn.
n.v. ).
Pigea filiformis DC. ex Ging. in DC. Prodr. 1 :307 (1824). Type: Port Jackson (holo: G-DC,
photo seen).
lonkUum fin forme (DC. ex Ging.) F. Muell., PI. Indig. Col. Viet. 1 :66 (1862)
Hybanthus filiformis (DC. ex Ging.) F. Muell., Fragm. 10:81 (1876).
Calceolaria filiformis (DC. ex Ging.) O. Ktze., Rev. Gen. PI. 1 :4I (1891).
loniclium linearioides PresI, Bot. Bemerk. 12 (1845), Type: " Habitat in Nova Hollandia ad
Port Jackson, collegit Sieber."' (holo: PR, n.v.).
Hybanthus tatei F. Muell. ex Tate. FI. Extratrop. S.A. 19 (1890). Type: Wilpena SA
(holo; MEL).
Calceolaria tatei (T. Muell. ex Tate) O. Ktze., Rev. Gen PI. 1 ;4I (1890).
Perennial herbs lo 60 cm tall. Leaves alternate (upper leaves j opposite),
linear to oblong. 5-90x0 -5-4 mm; stipules linear-lanceolate to triangular-
deltoid, 0-25-1 -5 mm long. Inflorescence axillary racemose, to 20 cm long.
Anterior petal 5-21 mm long. blue. Lateral petals of dissimilar pairs, mauve, 2
outer lanceolate acute, 2 inner broadly lanceolate-falcate, oblique, rounded
edge raised with several enlarged cells. Anthers: loculi appendages (if present)
narrow triangular, <0-5 mm, white or orange. Capsule 6 mm long; seeds 3-6
reticulate to reticulate-foveolate, blaek.
Distribution: This species is found in South Australia, S.E. Victoria, New South
Wales and eastern Queensland (Figure 6). 265 collections were seen.
gHpemW. Wallangarra, W. T./ortw 2617 (CANB); Mt. Barney slopes, Maepherson Ranges
15 Nov. 1952, E. F. Constable (NSW); Bribie Island and Pine River, Eaves (MEL)- Callabah’
S. L. Evera/ 31 16 (8R1). ’
New South Wales: Clarence River, 1869, Wdroy (MEL); near Tinda Creek, Putty Road 20
Feb. 1961,A/. E. P(t(7/(pv(CBG);inbushlandnearBundanoon, 29Nov. 1961 Ga«6a(GAUBA)
Khyber Pass, east of Rylstone, H. S. McKee 936 (SYD); Wollomombi, 17 Dec 1950 G L
Davis (AD); 2 miles W ^Karuah, 12 Oct., 1953, L. A. S. Johnson (NSW); Blac'kheath Nov'
1914, A. A. Hamilton GAS'N).
Victoria: Avon and Mitchell River, Dec. 1884, F. Mueller (MEL); Nitta Nilta Oct 1919
S. F. Cliton ( M EL).
South Australia: Wundinna, 7 Apr. 1936, E. H. Ising (AD); Summit of Mt. Woodroffe Mus-
grave Ranges, D. E. Symon 2663 (ADW). ’
234
Under lonidium monopetalum Roemer and Schultes gave a short diagnosis
and cited Viola monopetala as a manuscript name. The authors gave the origin
of the type specimen as “ Ex Anglia habiiit Romer in Herbario suo.” The
bulk of the specimens on which Roemer and Schultes descriptions were based
were housed at LZ and are now destroyed.
The name “ Viola monopctaki ” which was cited in synonymy by Roemer
and Schultes was used by Robert Brown both in his unpublished manuscript
and in his herbarium for the same species as that described by Roemer and
Schultes. It is possible that it was through Brown that Roemer and Schultes
obtained the type. This means that the collection of Robert Brown's housed
at the British Museum (Natural History) is a possible isotype. A photograph
of a plant collected at Port Jackson, on May 11, 1802 by Robert Brown has
been seen.
6. Hybanthus debilissimus T. Muell., Fragm. 1 1 ;4 (1878).
In silvis Karri-forests seciis flumen Shannon. (W.A.), F. Mueller, (holo: MEL; iso: AD).
Calceolaria dehiUssima (F. Muell.) O. Kuntze Rev. Gen. PI. 1 :4I (1891).
Perennial herb, procumbent or low-growing. Leaves alternate, sessile or
shortly petiolate, lanceolate. 5-25x2-5 mm, stipules not present (minute?).
Inflorescence of 1-3 flowers in axillary recemes, 2-3 cm long. Sepals narrow-
lanceolate, 2-4 mm long, acumimate, glabrous, deep-blue edged with white.
Anierior petal 5-12 mm long, blue. Lateral pends of dissimilar pairs, 2-3 mm
long, mauve, 2 outer petals narrow oblong, 1-2 nerved; 2 inner petals broadly
Figure 6^Distribution of Hybanthus monopetalus (Roem. et Schult.) Domin (•), H. debilis-
simus F. Muell. (x) and H. calycinus (DC. ex Ging.) F. Muell. (o).
235
oblong, 3-4 nerved. Stamens: paired loculi appendages (when present) narrow-
triangular, <0'5 mm long, white or orange. Capsule 5 mm long; seeds ca.
6. dark brown, pusticulate.
Distribution: This species is found in the Warren and Darling districts of
Western Australia (Figure 6). 26 collections were seen.
Western Australia: Stream banks on outskirts of Collie, A. R. Fairatt 760 (PERTH); near Pem-
berton, 18 Oct. 1919, C. G. Sargent (NSW); near Mt. Lindsay, 1879, Muir (MEL); Hamden,
Clarke (MEL); Margaret River, R. D. Royce 4683 (PERTH).
Mueller, in his original description, stated that there are small stipules
present at the base of the leaves, but none of the herbarium specimens (including
the type) shows these.
7. Hybanthus calycinus (DC. ex Ging.) F. Muell., Fragm. 10:81 (1876).
Pigea catycina DC. ex Ging. in DC., Prodr. 1:307 (1824). Type: Nouv. Holland: Cote
occidentale (holo: P, photo seen).
Pigea glauca Endl. in Endl. et al,, Enum. PI, Huegel 5 (1837). Type: Freemantle ad Swan-
River, Hiiget (holo: W).
Sotea caiycina (DC. ex Ging.) Spreng., Syst. Veg. 1 :804 (1825).
Vtamingia australasiaca De Vr. in Lehm., PI. Preiss. 1:399 (1845). Tvpe: Herb. Preiss,
No. 1449 (holo: W).
lonidium glaucum (Endl.) F. Muell., PI. Col. Viet. 1 :67 (1862).
tonidium catycinum (DC. ex Ging.) F. Muell., PI, Col. Viet. 1 :224 (1862).
Calceolaria caiycina (DC. ex Ging.) O. Ktze., Rev. Gen. PI. 1:41 (1891).
Perennial herb to 60 cm tall. Leaves alternate, sessile, linear, 2-4 -Sx
0 - 1-0-3 cm, glabrous or scabrous; stipules linear-lanceolate, 0-2-10 mm long,
glabrous i pubescent. Inflorescence a 5-to many-flowered axillary raceme,
ca. 20 cm long. Sepals lanceolate, 3-7 mm long, prominently 3-5 nerved,
glabrous or with ciliate margins, blue with white margins. Anterior petal
10-16 mm long, blue to purple. Lateral petals of dissimilar pairs, 3 -5-4 -5 mm
long, margins recurved, ciliate, blue to purple, prominent purple veining.
Stamens densely pilose, purple; paired loculi appendages narrow-triangular,
<0-5 mm long, orange. Capsule 6 mm long; seeds 3-6, ovoid to ellipsoidal,
2-5 mm long, black, prominently reticulate.
Distribution: This species is found in the coastal regions of the Irwin, Darling,
and Warren districts of Western Australia (Figure 6). 123 collections were
seen.
Western Australia: 45 miles NE. Perth, F. W. Went 65 (PERTH); Moresby Range Howatharra
A. C. Burns 3 (PERTH); 43 miles N. of Geraldton A. S. George 9182 (PERTH)- Cape Nat-
uraliste, E. M. Bennett 2269 (PERTH); Ogilvie, N. T. Burbidge 2142 (CANB); Leederville
1 Aug. 1897, R. Helms (WK\)-, King’s Park Perth, Dec. 1910,7. Sheath (NSW); Between Cham-
pion Bay and Port Gregory, Oct. 1877, F. Mueller (MEL): 18 miles N. of Bunburv 21 Oct
1962, M. E. Phillips (CBG).
Group 111; Flowers solitary
8. Hybanthus volubilis E. M. Bennett sp. nov. (Figure 7).
Herbci perennis volubilis. Folia altema, sessilia, vel linearia vel lanceolata herbacea-
stipulae lineares. Flores axillares, solitarii, violacei. Sepah lineari-lanceolata acuta re-
curvata. Petalum anticum manifeste calcaratum, album venis atroviolaceis, petala lateralia
oblonga. Loculi antherarum appendicibus anguste-triangularibus. Ovarium ovoideum vel
globosum, glabrum, ovulis 6.
Twining perennial herb. Leaves alternate, sessile, linear-lanceolate
10-18x2-4 mm, occasionally basal leaves 16-20x5-6 mm chartaceous glabrous’
stipules linear, glabrous, 0-25 mm long. Flowers axillary, solitary Sepals
lanceolate to linear, acute, 2-2 -5 mm long, recurved, green to purple. Anterior
petal 6-8 mm long, lamina white with mauve veining. Lateral petals oblong
apices truncate, recurved, 1-5-2 -5 mm long, blue-mauve with darker veinine
236
Figure 7 — Hybanthus volubilis sp. nov. A — Inflorescence. B — Posterior surface of anterior
anther. C — Anterior surface of anterior anther. D — Posterior anther. E — Posterior
surface of inner petal. F — Anterior surface of inner petal. G — Outer petal. H — Sepal,
J — Ovary. K — Leaf. L — Anterior petal. M — Capsule, a = apex recurved; s = spur; r =
raised ridges : 1 = anterior petal, white with mauve markings.
237
Anthers green-yellow; loculi appendages narrow-triangular, 0-25-0-5 mm long,
orange-yellow. Ovary ovoid to globose, glabrous, ovules 6. Capsule 5 mm
long; seeds 1-6 per capsule, ovoid-ellipsoidal, ca. 2-5 mm long, smooth, mottled
pale and dark brown.
Type: 8 miles E. of Margaret River townsite on banks of Margaret River, W.A., 17 Nov. 1968,
E. M. Bennett 2815 (holo: PERTH).
Distribution: This species is found in the Warren district of Western Australia
near the townsite of Margaret River (Figure 8).
Western Australia: Rosa Brook, R. D. Royce 2803 (PERTH); on banks of Margaret River,
Osmington, R. D. Rovce 2464 (PERTH); Osmington, Margaret River district, R. D. Royce
4621 (PERTH).
9. Hybanthiis vernonii (F. Muell.) F. Mitell., Fragm. 10;81 (1876).
lonidinni vernonii F. Muell., PI. Indig, Col. Viet. 1:223 (1862). Type: Twofold Bay, N.S.W.,
F. Mueller (syn; MEL, K, photo); Genoa River, Vic., Sept. I860, F. Mueller (syn.: BM,
photo seen).
Calceolaria vernonii (F. Muell.) O. Kuntze, Rev. Gen. PI. 1;4I (1891).
Hyhanthus enneaspermus (L.) F. Muell. var vernonii (¥ . Muell.) Domin, Biblioth. Bot. 89:982
(1928).
Perennial herb or shrub to 90 cm tall. Leaves alternate, sessile, basal
leaves usually lanceolate, upper leaves linear 5-45x1-4 mm, stipules ca. 0-5 mm
long, linear. Flowers solitary in axils of upper leaves. Sepals lanceolate, 1-5
to 5 mm long, glabrous or outer surface scabrous, green to dark blue, margins
238
■±. white. Anterior petal 6-13 mm long, blue to mauve. Lateral petals of
dissimilar pairs, I -5-3 mm long, blue to mauve, 2 outer petals narrow-oblong.
1-nerved, 2 inner petals broadly oblong, several nerved. Stamens: anterior
anthers densely pubescent on surface, posterior 3 i shortly pubescent; loculi
appendages white, narrow-triangular, <0-5 mm long. Capsule 6-8 mm long;
seeds 3-6, ovoid-ellipsoidal, ca. 2-5 mm long, brown, smooth to lineolate.
Key to subspecies
Leaves and stems glabrous. Upper leaves linear, lower usually lanceolate subsp. vernonii
Leaves and stems scabrous. Leaves lanceolate, oblanceolate or tricuspidate subsp. scaber
subsp. vernonii
Leaves and stems glabrous. Lower leaves much broader than upper
leaves, linear to lanceolate, 5-45x1-4 mm.
Distribution: This subspecies is found along the Central and South coast of
of southern New South Wales and Eastern Victoria (Figure 9).
57 collections were seen.
New South Wales: Kiola Stale Forest, South Coast, 26 Oct. 1966, M. Evans (CANS); Jervis
Bay, 28 Sept. 1961, Gauba (GAUBA); National Park N.E. Sector, 16 Aug. 1927, O. D. Evans
(SYD); Middle Harbour Sydney, Sept. 1909, J. B. Cletand (AD); Green Cape Track, 8 Oct.
1961, M. E. Phillips (CBG); Port Jackson, W. Kmio// (MEL); Gynea, R. H. Goode 456 (NSW).
Victoria Gippsland, Point Ricardo, L. B. Mini- 3783 (MEL); near mouth of Snowy River.
E. Pescott 14 (MEL).
Figure 9 — Distribution of Hybanthus vernonii (F. Muell.) F. Muell. subsp. vernonii (•) and
subsp. scaber (o).
239
subsp. scaber E. M. Bennett subsp. nov.
Cmiles costis scabris, Folia 5- 1 5x1 -5-2 -5 mm, vel lanceolata vel oblanceolata yel
triciispidata, pilis in paginis ambabus sparsis, praesertim abundioribus in venis et in margin-
ibus recurvis.
Sleim scabrous along ribs. Leaves 5-15x1 ■5-2-5 mm varying from lance-
olate to oblanceolate to tricuspidate, with scattered hairs on both surfaces,
especially abundant along veins and recurved margins.
Type: Stockout Creek, Coledale Road, N.S.W., Oct. 1909, J. L. Boorman (holo: NSW 97630).
Distribution: This species is found along the east coast of New South Wales
(Figure 9). 14 collections were seen.
New South Wales: Track to Green Cape, 8 Oct. 1961. M. E. Phillips (CBG); Bundanoon, 21
Apr. 1934, L. R. F. and J. W. V. (SYD); Clyde district, Dec. 1884, W. Baeuerlen (MEL);
Stockyard Gully near Copmanhurst, 23 Jul. 1967, K. Grieve (NSW); near Turpentine Nowra-
Braidwood, 15 Jul. 1934, F. A. Rodway (NSW).
10 . Hybanthus epacroides (C. A. Gardn.) Melch., Nat. Pflanzenfam. Ed.
2 -21 :360 (1925).
Jonidiuni epacroides C. A. Gardn., J. and Proc. Roy. Soc. W. Austral. 9:35 (1923). Type:
near Mt, Marshall, half a mile south of Bencubbin Station, 2 June 1922, C. A. Gardner i696
(holo: PERTH; iso: MEL).
Shrub to 40 cm high, spinescent, lateral branches divaricate. Leaves
clustered at nodes, sometimes alternate, linear to spathulate, 2-7 x 0-5-1 0 mm,
apex recurved; stipules linear to lanceolate, ±0-5 mm long, glabrous except
for few hairs towards base. Flowers axillary or arising from middle of leaf
cluster, solitary. Sepals ovate-lanceolate, ca. 1-5 mm long, glabrous or
ciliate, pale blue to white. Anterior petal 3-5 mm long, pale blue. Lateral
petals of dissimilar pairs, oblong 1-5-3 mm long, apices truncate, recurved,
pale blue to white, veining purple. Anthers purple, ± pubescent; loculi ap-
pendages absent. Capsule 5 mm long; seeds 1-2, ovoid-ellipsoidal, ca. 3 mm
long, smooth, mottled pale and dark brown.
Distribution: This species is found in the Coolgardie and Eyre districts of
Western Australia (Figure 8). 22 collections were seen.
Western Australia: Lake Cronin, Forrestania, 13 June 1929, C. A. Gardner (PERTH); 146
miles Great Eastern Highway, E. M. Scrymgeour 689 (PERTH); Nangeenan, F. Stoward 3
(NSW); 2 miles West of Norseman, 17 September 1962, M. E. Phillips (GBG).
11 . Hybanthus bilobus C. A. Gardn., J. Roy. Soc. W. Austral, 19:86 (1933).
Type: Hamersley River, near Eyre Range, 23 Sept. 1925, C. A. Gardner 1886 (holo: PERTH)
H. floribundus (Lindl.) F. Muell, var. minutifoUa F. Muell, Fragm. 10:82 (1876) Type-
Crescit a Phillips River usque Cape Arid, Maxwell (holo; MEL).
Shrub to 20 cm tall, lateral branches divergent forming a dense intricate
network. Leaves scattered, usually crowded in small clusters at the nodes,
sessile, obdeltoid, I -2x0 -5- 1 -5 mm, truncate, appearing bilobed as a result of
the recurved central portion of the apex; stipules linear, ±0-5 mm long, densely
hairy in lower half. Flowers axillary or arising from middle of leaf cluster,
solitary. Sepals lanceolate to oblancolate, 1-5-2 -5 mm long, densely hirsute
along margins and veins. Anterior petal 3-4 mm long, pale blue. Lateral
petals of dissimilar pairs, oblong, 2-3 mm long, pale blue, veining purple
Anthers blue or green, pubescent, loculi appendages not present.
Distribution: This species is found in the Eyre and Coolgardie Districts of
Western Australia (Figure 8). 10 collections were seen.
Mungilginup River, ± 50 miles east of Esperance, 8 Sept 1963 / H Willis
(MEL); 544 mile peg Norseman-Esperance road, E. M. Bennett 2168 (PERTH)- 5 miles S nf
Salmon Gums, 10 Aug. 1951, N. H. Brittan (UWA). ’
240
References
(Works cited in introductory pages)
Airy Shaw, H. K. (1966) Revised 7th ed. of J. C. Willis, A Dictionary of the Flowering
Plants and Ferns. Cambridge University Press, England.
Dandy, J. E. (1969) — Nomina Conservanda Proposita. “Calceolaria”. Taxon 18:469.
Darlington, C. D., and Wylie, A. P. (1955)— Chromosome Atlas of Flowering Plants.
G. Allen and Unwin, London.
Diels, L. (1904) — Fragmenta Phytographiae Australiae occidentalis. Bot. Jb. 35:56.
Hutchinson, J. (1967) — The Genera of Flowering Plants. Vol. II. Clarendon Press, Oxford.
Lawrence, F. H. M. (1951)— Taxonomy of Vascular Plants. Macmillan Co., New York.
Martin, A. C. (1946) — The comparative internal morphology of seeds. Am. Midi. Nat.
36:513-660.
Melchior, H. (1925)— In Engler, A. Die Naturlichen Pflanzenfamilien. Duncker and Hum-
blot, Berlin.
Mueller, F. (1876)— Fragmenta Phvtographiae Australiae. 10:81-82.
Ornduff, R. (Ed.) (1967)-Index for Plant Chromosome Numbers for 1965. Regnum Veg.
Vol. 50.
Raju, M. V. S. (1958)— Seed development and fruit development in lonidium suffhilicostim
Ging. Phytomorph. 8:218-224.
Rickett, H. W. (1944)— The classifications of inflorescences. Bot. Rev. 10:187-231.
Rickett, H. W. (1955) — Materials for a dictionary of botanical terms— 111 Inflorescences.
Bull. Torrey Bot. Club 82:419-445.
St. Hilaire (1824, n.v.)— Mem. Mus. Par. 11.
Schulze, G. K. (1936) — Morphologisch-systematische Studien fiber die Gattung Hybanthus
mit besonderer Berficksichtigung der sudamerikanischen Arten. Bot. Jb. 67:437-492.
241
Figure 1— A, Q^Eucatyplus pendens sp, nov. A— Buds (Brooker 1853) B— Fruit (Brooker
F- (Middle Mt, Barren Se?
2976). D Fruit (Gbi drier 2976). E — Fruit (cultivated Hamel nurserv, FRI 15797)
242
Four new taxa of Eucalyptus from Western Australia
By M. I. H. Brooker*
Abstract
Four taxa of Eucalyptus are described — E. pendens from the Badgingarra
district, E. acies from the Fitzgerald River Reserve, E. loxophleha subsp. gratiae
from the Lake Grace district, and E. conglohata subsp. fraseri from the country
north of Esperance and east of Norseman.
Eucalyptus pendens M. I. H. Brooker sp. nov. (Figures I A, B, 2)
Eucalypio sepulcrali F, Muell. aflfinis a qua alabastris minus elongatis et fructibus minor-
ibus non-urceolatis ditfert.
Frute.x " mallee ” 2-5 m altus ramificatione laxa demum pendente. Ramuli juvenes
quadrangulares cortice laevi atro-rubra nitenti glauca. Ramuli maturi et caules cortice
laevi cinereo-erubescenti. Glandulae oleorae in cortice et in medulla nullae. Lignotuberum
formans.
Cotyledones reniformes I -4-3 Ox I • l-l -5 cm, triplinerves, supra virides, infra purpureae.
Folia planiulae sessilia, decussata, elliptica vel ovato-oblonga undulatescentia, 2-5 x 1^
cm. Axis plantulae laevis. Folia intermedia breviter petiolata. decussata, late-elliptica,
valde undulata, usque ad 16x8 cm. Folia matura petiolata, in gemma decussata, demum ab
intranodiis separata, lanceolata, 6-12 x 1-2 cm costa in pagina una vel utrinque impressa,
dense reticulata, sine glandulis oleosis manifestis, viridia, concoloria. Petioli complanati.
2-3 cm longi.
Inflorescentiae axillares, 7-1 1( l3)-florae. Pedunculi erecli, validi, in sectione trans-
versal! ovales vel complanati ad apices laliores, I •5-4-0 cm longi. Involucrum 6-bracteatum,
bracteis 5-6 mm longis pro longitudine maxima connatis, demum deciduis. Inflorescentiae
bracteatae in axillis foliorum dum inflorescentiae maturae expansis in ramulis vetioribus
defoliatis praesentes. Alabastra clavata sine cicatricibus abscissis, 9-1 1 x 6-8 mm, in pedicellos
complanatos vel quadrangulares 3-8 mm longos angustata. Operculum hemisphaericum
interdum umbonaium, vel conicum, verrucosum inlerdum radiatim 2-3-costatum, hypan-
thium aequans vel brevius. Stamina omnia fertilia. Filamenta alba, in alabastro radiale
inflexa, eglandulosa. acuminata. Antherae dorsifixae versatiles, reriiformi-orbiculares,
ab rimis obliquis, ovalibus, non-confluentibus dehiscentes. Gians terminalis. Stylus gland-
ulosus. interdum ad apicem Hexus. Ovarium 4-5-loculare. in hypanthio profunde impressum.
Oviila verticaliter 2-seriata.
Fructus ovoideus vel cylindricus, truncalus, I •3-2-0 x I -2-2-0 cm, rugosus vel parum
costatus. Discus obliquus ad verticalis. Valvae 4-5, profunde inclusae.
Semina 4-5 x 3-4 mm, latere dorsali rotundata, ventral! cum porcis aliquot ad hilum ad-
scendentibus, nigra. Ovulodia angularia, quam semina minora, rufa.
Type - At 125 mile peg north of Perth on highwav between Gingin and Badgingarra, Western
Australia (30°14'S, 115 28'E), 23 .luly 1969, M. I. H. Brooker 1949. (holo: PERTH; iso:
PERTH, FRl, K).
Allied to E. sepuicnilis F. Muell. but differing in the less elongate buds
and in the fruit which are smaller and not urceolate.
A mallee 2-5 m tall with a light, finally drooping canopy. New hranchlets
quadrangular, bark smooth, dark red, shining, glaucous. Older branchlets
and main stems with pinkish-grey smooth bark. Oil glands absent in both
bark and pith. Capable of forming lignotubers.
Cotyledons reniform, 1 -4-3-0 x 1-1-1 -5 cm, triplinerved, green above,
purple beneath.
Seedling leaves sessile, decussate, elliptical or ovate-oblong, becoming
undulate, 2-5 x 1-4 cm. Seedling axis smooth. Intermediate leaves shortly
Forestry and Timber Bureau, Canberra, A.C.T. 2600
243
petiolate, decussate, broadly elliptical, markedly undulate, up to 16 x 8 cm.
Adult leaves petiolate, decussate in bud, finally separated on the axis by intra-
nodes, lanceolate, 6-12 x 1-2 cm the midrib impressed on one or both sides
densely reticulate, without apparent oil glands, green, concolorous. Petioles
flat, 2-3 cm long.
Inflorescences axillary, of 7-1 1 (13) buds. Peduncles erect, stout, oval or
flattened in transverse section, widening at the tip 1 •5-4 0 cm long. Involucre
of six bracts, 5-6 mm long, fused for most of their length, finally deciduous.
Bracteate inflorescences occuring in leaf axils while mature expanded inflores-
cences occur on older branchlets from which the leaves have fallen. Unopened
buds clavate, without an abscission scar, 9-1 1 x 6-8 mm tapering into the pedi-
244
cels which may be flattened or quadrangular, 3-8 mm long. Operculum
hemispherical and sometimes umbonate, or conical, verrucose and sometimes
with 2-3 radial ribs, equal to or shorter than the hypanthium. Stamens all
fertile. Filaments white and indexed radially in the bud, non-glandular,
acuminate. Anthers dorsifixed versatile, reniform-orbicular, opening by
oblique, oval, nonconfluent slits. Gland terminal. Style glandular, some-
times bent at the top. Ovary 4-5 locular, deeply sunk in the hyanthium.
Ovules in two vertical rows.
Fruit ovoid or cylindrical, truncate, 1 ■3-2 0 x I •2-2 0 cm, wrinkled or
shallowly ribbed. Disc oblique to vertical. Valves 4-5, deeply enclosed.
Seed 4-5 x 3-4 mm, dorsal side rounded, ventral side with several ridges
ascending to the hilum, black. C/jq//’ angular, smaller than seed, red-brown.
Distribution: Western Australia, near west coast in the region of the Hill River
120 to 150 miles north of Perth (Figure 2).
Other collections'. 1 19'6 miles N of Perth on Gingin-Badgingarra highway (30 18'S, 1 15°30'E),
14 May 1969, A. S. George 9312, 9314, 9315 (PERTH); 120 miles on same highway (30°I8'S,
I I5°30'E), 6 June 1969, Brooker 1850, 1852 (PERTH); 127-5 miles on same highway (30" I2'S,
115 27'E), 6 June 1969, Brooker 1853 (PERTH, AD, GAUBA, CANB, ERE K, MEL, NSW);
151 mile peg, Badgingarra-Eneabba road (30 02'S, II5°20'E), 30 Aug. 1969, A. S. George
9624 (PERTH).
Flowering period: July-August,
E. pendens is a remarkable species closely related to and resembling E.
sepulcralis but differing in bud and fruit morphology (Figure 1). The two
species occur approximately 350 miles apart and like some other Western Aus-
stralian eucalypts they are notable for their restricted occurrence. £. sepul-
cralis occupies a few sites on sandy foothills of the ranges between the Fitz-
gerald River and Hopetoun on the south coast, E. pendens occurs on lateritic
sandhills and has been collected only at the localities cited above. The species
occurs as isolated groups of slender mallees whose stems are only a few cm in
diameter and which emerge above low. dense, sclerophyllous shrubs. The
canopy is thin and always pendulous. There are no other Eucalyptus species
associated with it in the field.
Eucalyptus acies M. I. H. Brooker, sp. nov. (Figures 3 A-C, 4)
Frutex “mallee" I •0-2-5 m altus, vulgo pluricaulis, cortice laeve. /famw// juvenes com-
planali, quadrangulares. Glandulae oleosae in cortice et in medulla nullae.
Cotviedones emarginatae, reniformes, I •7-2-2 x I ■5-2 0 cm, supra atrovirides, infra
atromalvinae. Folia planudae sessilia, decussata, elliptica, 5-7 x 3-5 cm, parum discolona
Axis plantulae verrucosus, glandulosus. Folia matiira opposita vel sub-opposita, lanceolata
ad latelanceolata, symmetrica ad parum falcata, acuminata, coriacea, (8-5) 9 0-I3-5 (14-5) x
2-0-3-5 cm, viridia, concoloria, Petioli complanati (1-5) 2;0-3-0 cm longi. Costa mediana
distincta, infra prominentior. paribus multis venarum lateralium ab costa mediana sub angulo
(40 ) 45"-55 (60 ) prodeuntibus. Vena intramarginalis ab margine 1-2 mm distans.
Inftoresceiitiae axillares 7-florae. Pedunculi complanati, biconvexi, ad apices latiores.
saepe reflexi, (1-3) I ■7-2-3 (2-6) cm longi. Gemma inflorescentiae inapertae rotundata, ab
bracteis 4 inclusa.
Atahaslra clavata sine cicatricibus abscissis, cum pedicellis I -5-2 - 1 x 0-7-1 ■ I cm longa.
Hvpanthium parum angulare, manifeste costatum. quam operculum vulgo multo longius,
0-9-1 -4 cm longum, in pedicellum angustatum. Operculum depresso-hemisphaericurn,
laeve vel parum costatum, saepe breviter rostratum. Stamina omnia fertilia. Filamenta in
alabastro radiale intlexa, eglandulosa. acuminata. Amherae dorsifixae, versatiles, reniformi-
orbiculares, ab rimis longitudinalibus obliquis dehiscentes. Gians terminalis. Stylus con-
icus, rectus, alabastro supra filamenta elevatus. Stigma inconspicuum. Ovarium quam
hypanthium multo breviore, 3(4)-loculare. Ovula verticaliter 2-seriata.
Fructus hemisphaerico-campanulaia costibus prominentibus aliquot, I 1-1 -6 x 1 -2-1 -6
cm, in pedicello brevi in sectione transversali triangulari 3-8 mm longo attenuala. Annulus
calycinus angustus, planus, manifestus; discus latus, conspicuus, planus vel convexus; valvae
3(4), rudimentales, cum disco aequatae.
245
Figure 'i—A-C —Eucalyptus acies sp. nov. A- -Buds (Brooker 2723)
2723). C— Seedling stem (cultivated FRI nursery). D— Eucalyptus
Seedling stem (cultivated FRI nursery).
B — Fruit (Brooker
preissiaua Schau.
246
Semina angulata porcis parum alatis, 2-3 x 2 mm, atro-cinerea vel nigra, aliquar.tum
nilentia.
Type: On Woolbernup, a rocky hill in the Fitzgerald River Reserve between Bremer Bay and
hiopetoun Western Australia (34 02'S. 119 4rE), 4 Aug. 1970, M. I. H. Brooker 2725 (hole:
PERTH iso: PERTH. GAUBA, FRI, K, MEL, NSW).
A malice shrub 1-0-2-5 m tall, usually several-stemmed, with smooth
bark. Young hranchlets flattened and quadrangular. Oil glands absent in
both bark and pith.
Cotyledons emarginatc, reniform, I ■ 7-2-2 x I •5-2-0 cni dark green above,
deep mauve below. Seedling leaves sessile, decussate, elliptical, 5-7 x 3-5 cm
slightly discolorous. Seedling axis verrucose, glandular. Adult leaves opposite
to sub-opposite, lanceolate to broad-lanceolate, symmetrical to slightly falcate,
acuminate, coriaceous, (8-5) 9 0-13-5 (14-5) x 2-0-3-5 cm green, concolorous.
247
Petioles flat (1-5) 2 0-3 0 cm long. Midrib distinct, more prominent below,
with numerous pairs of primary lateral veins at an angle of (40°) 45°-55° (60°),
Intramarginal vein 1-2 mm from the margin.
Inflorescences axillary, of 7 flowers. Peduncles flattened, biconvex, and
broader at the top, often reflexed (1-3) l -7-2'3 (2-6) cm long. Unopened
inflorescence bud rounded, enclosed by 4 bracts.
Unopened buds clavate without abscission scars I -5-2 -I x 0-7-1 1 cm
including the pedicel. Hypanthium slightly angular, with prominent ridges,
usually much longer than the operculum, 0-9-1 -4 cm, tapering into the pedicel.
Operculum depressed-hemispherical, smooth or shallowly ribbed and often
shortly beaked. Stamens all fertile. Filaments indexed radially in the bud,
non-glandular, acuminate. Anthers dorsifixed, versatile, reniform-orbicular,
opening by longitudinal-oblique slits. Gland terminal. Style conical, erect,
raised above filaments in the unopened bud. Stigma inconspicuous. Ovary
much shorter than the hypanthium, loculi 3 (4). Ovules in 2 vertical rows.
Fruit hemispherical-campanulate, with several prominent ridges, 1 - 1-1 - 6 x
1 -2-1 - 6 cm, tapering into a short pedicel, triangular in cross-section, 3-8 mm
long. Calycine ring narrow, flat and well defined; disc wide, conspicuous,
level to convex; valves 3(4), rudimentary, more or less level.
Seeds angular, with slightly winged ridges 2-3 x 2 mm, dark grey or black,
rather shiny. Hilum area conspicuous. Chajf light red-brown, angular,
smaller than the seed.
Distribution: Western Australia, south coast between Bremer Bay and Hope-
toun (Figure 4).
Other collections. Thumb Peak Range (34°02'S, I19°43'E), 31 Oct. 1965, A. S. George 715o
(PERTH); Thumb Peak (summit) (34’02'S, 119°43'E), 23 Oct. 1970, R. D. Royce (PERTH);
Middle Mt. Barren (34°03'S, n9°4rE), 16 July 1970, A. S. George 10088 (PERTH); Wool-
bernup (34°02'S, 1 19”41'E), 4 Aug. 1970, M. /. H. Brooker 2723 (PERTH).
Flowering period: September-November.
E. acies is a species of notably restricted occurrence. Its distribution is
known only from the above localities which are themselves only a few miles
apart. It has not been collected on the intervening plains. The species is
therefore likely to be an endemic in the Fitzgerald River Reserve.
The natural affinity of E. acies is not clear. It belongs in the Renantherae
and is probably closest to £. preissiana which it somewhat resembles in habit
and gross morphology. It differs from E. preissiana in having 7 buds to the in-
florescence, ribbed hypanthia, filaments which are apparently non-glandular,
and glandular-verrucose seedling stems compared with the stellate hairy stems
of E. preissiana (Figure 3 C, D).
The Series Preissianae of Pryor and Johnson (1971) consists of three renan-
therous species, E. megacarpa, E. preissiana and E. coronata which were in-
correctly included by Blakely in the Section Macrantherae. Each has 3 buds to
the inflorescence and glandular filaments and it may be incorrect to classify
E. acies with equal status w'ithin this Series. As E. acies does not resemble other
known species 1 suggest it tentatively be placed in the Series Preissianae or be
considered as a related, as yet monotypic, Series.
Eucalyptus loxophleba Benth. subsp. gratiae M. I. H. Brooker subsp. nov.
(Figures 5, 6).
• subspecie typica foliis, alabastris, tructibusque majoribus differt. Cortex laevis
mtjdus. RW; plerumque glauci Ro/w 7-1 1 x I -2 - 5 cm, nitida. Pedunculi complanafi,
Indusis^'^* -4/a6o.s//-a 12-15 x 4-5 mm. Fructus obconica usque ad 12 x 9 mm pedicellis
248
Type: 0-5 mile west of Burngup, Western Australia (33°OI'S, 1 18°4rE), 3 Nov. 1969, M. /. H.
Brooker 2273 (holo: PERTH).
Differs from the typical subspecies in the larger leaves, buds and fruit.
Bark smooth, shining. Branchlets usually glaucous. Leaves l-\ \ x l-2'5 cm,
glossy. Peduncles flattened, broad, glaucous. Buds 12-15 x 4-5 mm. Fruit
obconical, up to 12x9 mm, including the pedicel.
Distrihution : Western Australia, south-west, between Dumbleyung and Lake
King (Figure 6).
Other collections: 4 miles east of Dumbleyung (33 I6'S, 117 46'E). 29 Oct. 1962, M. E.
Phillips (CBG 023097); 12 miles east of Newdegate (33 06'S, 119 I4'E). 30 Oct. 1962, M. E.
Phillips (CBG 023090); 15-5 miles west of Lake King (33 06'S, 119'23'E) 17 Mar. 1967,
G. M. Chippendale 225 (FRI); 25-7 miles north east of Lake Grace township (32'’52'S,
I I8°39'E), 17 Mar. 1967, G. M. Chippendale 230, (FRI); 25 miles west of Lake King (33'’06'S,
119°I4'E), May 1969, B. A. Rockel (FRI 18598); 10 miles west of Lake Grace (33°06'S,
!18°16'E), 3 Nov. 1969, M. I. H. Brooker 2268, 2269 (PERTH, FRI, NSW); 0 ■5 mile east
of Cargonocking Hill (32°49'S, 118 '04'E), 13 July 1970. M. I. H. Brooker 2652 (PERTH).
Flowering period: Jiine-Octoher.
Figure 5 — Encalyplus loxophleha subsp. gratiae subsp. nov. Fruit (25-7 miles north east of
Lake Grace, Chippendale 230).
E. loxophleha is a widespread species in south west Western Australia in
the 10-20 in, rainfall region. Published by Bentham in 1867, its status was
doubted by Mueller (1884) who considered it may only have been a tree form
of “ E. foecunda " (£. oraria L. A. S. Johnson). This suggestion was adopted
by Lueitmann (1898) who indicated it was a variety of “ E. foecunda " and by
Maiden (1909) whose comments on the taxonomy of £. loxophleda were con-
fined to quotations from Mueller and Luehmann. Blakely restored £. loxo-
phleha to specific status (1934) and it has since been regarded as a distinct
species. Blakely, however, seems to have assumed its affinity with “ £. foe-
cunda " (£. oraria) by placing these two species next to each other in his classi-
fication. This anomaly w'as recognised by Pryor and Johnson (1971) who
place £. loxophleha in the Section Bisectaria and £. oraria in the Section Dum-
aria. The species of these two Sections are separable on cotyledon shape.
249
97093 — ( 2 )
//S"
Figure 6 — Distribution of Eucalyptus loxophleba Benth. subsp. gratiae subsp. nov.
E. loxophleba is a unique species and when a complete specimen is available
it is easily recognizable. It is characterised by bisected cotyledons; glandular
pith; prominent leaf venation with the intramarginal nerve well in; thick,
inflected filaments in several series, the outer ones long, elbowed in the bud and
strongly spreading in the flower, the inner ones short and erect in the flower,
some filaments strongly sculptured at the base, all narrowing suddenly to the
attachment with the connective; style constricted at the base.
While the above characters are invariable the species is notoriously variable
in habit and bark. Typical specimens as described in Bentham are trees with
“ rough ash grey fibrous bark This description would suit the species as
known in the higher rainfall areas, e.g., at York whence comes its common
name. However, the species appears to vary clinally from the higher rainfall
to the lower rainfall areas of distribution in that the tree habit is lost to the
north and east and also the amount of rough bark becomes less and less until
in the dry lands (c.g. Carrabin) £. loxophleba is a smooth-barked mallee. Such
a dine form is probably what is represented by £. loxophleba var. fniticosa
which Bentham described as a shrub from the Murchison River.
By contrast, subsp. gratiae is a distinctive form which occupies a small
area of distribution between Dumbleyung and Lake King and the sub-specific
name is derived from Lake Grace where it is prominent. It is notable in the
250
field for its fairly dense rounded canopy of bright green leaves, the smooth
trunks and the bright green-leaved seedlings.
Eucalyptus conglobata (R. Br. ex Benth.) Maiden subsp. fraseri M. I. H. Brooker
subsp. nov. (Figures 7, 8).
A subspecie typica habitu et alabastrorum forma differt. Arbor erecta ad 20 m alta cortice
laevi alba vel cinerea (interdum per 0-5 m e basi fibrosa). Alabastra variabilia, sessilia ad
breviter pedicellata, operculis pyramidalibus, rostratis vel obtuse conicis, costatis.
Type: 14 miles west of Balladonia, Western Australia, 14 Feb. 1970, M. I. H. Brooker 2472.
(holo: PERTH; iso: PERTH, AD, GAUBA, BRI, CANB, FRI, K, MEL, NSW),
Differs from the typical subspecies in habit and the form of the buds. An
erect tree to 20 m tall with smooth white to whitish-grey bark (or sometimes
with 0-5 m of basal rough bark). Buds variable, sessile to shortly pedicellate,
with pyramidal, rostrate or obtusely conical, ribbed opercula.
Distribution: Western Australia, north of Esperance and east of Norseman.
(Figure 8).
Other collections: 2 miles south from Beete Sidina (32'’44'S. I2I°32'E), 7 Nov. 1953, C. A.
Gardner 11161 (PERTH); near Balladonia (32 27'S, 123 51'E), 9 Sept. 1962, M. E. Phillips
(CBG 021970); east side of pass in Fraser Range (32 02'S, 122 52'E). 18 Oct. 1966, A. S
George 8597 (PERTH); 67-4 miles east of Norseman (32 02'S, 122 52'E), 12 Mar. 1967,
G. M. Chippendale 158 (FRI); between 497 and 498 mile peg on Norseman-Esperance Road
(32 04'S, 122 29'E). April 1969, B. ,-(. Rockel (FRI 18590); 8 miles north of Salmon Gums
(32 52'S, 121 36'E). 15 Feb. 1970, M. I. H. Brooker 2495 (PERTH); 6 miles east of Scaddan
(33 56'S, 121 50'E), 8 Aug. 1970, M. I. H. Brooker 2772 (PERTH); between 497 and 498
mile peg on Norseman-Esperance Road (32 04 'S, 122°29'E), 13 Sept. 1970, J. Baker 61 (FRI).
Flowering period: February.
Eucalyptus dittnosa var. conglobata R. Br. ex Benth, was published by
Bentham (1867) who cited two South Australian localities, viz., “ Port Lin-
coln ” (Wilhelmi) and “ South Coast " (R. Brown). Maiden (1909) published
it as a variety of E. incrassata Labill. and subsequently raised it to E. conglobata
(1924). In doing so he inadvisedly chose a later collection from Port Lincoln
as the type. It is now' known that the Robert Brown specimen at least is
extant and is lodged in the Herbarium of the Royal Botanic Gardens, Kevv
(G. Chippendale pers. comm.).
In the original description of var. conglobata Bentham (loc. cit.) did not
(mention habit though for E. dittnosa the habit is given as “ a shrub or small
tree Bentham's description of the flowers of var. conglobata as being
“closely sessile, the calyx-tube shorter than broad, angular, and operculum
conical ” fits the Port Lincoln specimens but is inadequate to eimbrace all the
forms which are now known to occur in the wide distribution from Western
Australia to Encounter Bay in South Australia.
One such form is the striking smooth-barked inland tree with the variable
bud form (subsp. fraseri) compared with the coastal mallee with the strictly
closely sessile buds which is typical of the form in the Port Lincoln district and
the south coast of Western Australia (subsp. conglobata). Maiden (1924)
reported trees of E. conglobata on Boston Island near Port Lincoln which were
“ 50 feet high ". Further collecting is required to sort out the relationship
of these trees to subsp. conglobata and subsp. fraseri.
Pryor and Johnson (1971) accepted E. conglobata as a distinct species to
include E. anceps R. Br. ex Maiden (Blakely) which is given anticipated sub-
specific status. In this scheme subsp. fraseri should be given taxonomic status
equal to both “ conglobata ” and “ anceps ”.
251
Figure 1— Eucalyptus conghbatu (R. Br. ex Benth.) Maiden subip. fraseri subsp. nov. A—
Bark (Brooker 2495). B Buds (Brooker 2472). C — Fruit (Brooker 2472).
252
C£NT1fcgTRgS
ROCK
)ailadcnia Hs.
‘Dowak
iSalrnon
MiLESIt
4C MILES
Figure 8— Distribution of Eucalyptus conglohala (R. Br. ex Benth.) Maiden subsp. frateri
subsp. nov.
Acknowledgements
1 would like to thank Mr. A. Popplewell for drawing my attention to
E. conglohata subsp. fraseri, and Mr. A. S. George for drawing my attention
to E. pendens and E. acies and for all the latin descriptions. Thanks are also
due to Mr. J. Baker for assistance with the description of E. acies, and to Mr.
N. Hall and Mr. A. Edward for the photographs.
References
Bentham, G. (1867) Flora Australiensis, Vol. 3 (L. Reeve & Co., London).
Blakely, W. F. (1934)- A Key to the Eucaiypts. (The Worker Trustees, Sydney).
LuEitMANN, J. G. (1898) Proc. Australasian Assoc. Adv. Sc. 7 (Sydney).
Maiden, J. H. (1909)— A Critical Revision of the Genus Eucalyptus Vol. I (Govt. Printer,
Sydney),
Maidf.n, j. H. (1924)- A Critical Revision of the Genus Eucalyptus Vol. 6 (Govt. Printer,
Sydney).
Mueller, F. (1879-1884) -Eucalyptographia (Govt. Printer, Melbourne).
Pryor, L. D. and Johnson, L. A. S. (1971) A Classification of the Eucaiypts (Australian
National University, Canberra).
253
Studies in the genus Acacia — 1
By B. R. Maslin
Abstract
Two species and one subspecies of Acacia are described — A. megacephala
sp. nov., A. echinata sp. nov. and A. moirii E. Pritzel subsp. recurvistipiila
subsp. nov. All three taxa belong to Bentham's series Pulchellae.
Introduction
This is the first in a series of papers which will deal with the taxonomy
of Acacia species primarily from Western Australia. Some of the morpho-
logical terms used in this and subsequent articles require brief explanation.
1. Receptacle: that portion of the inflorescence axis on which the flowers
are borne.
2. Bracteoles: small bracts positioned on the receptacle, each of which
subtends one flower.
3. Terminal seta: Bentham, Journ. Bot. (Hooker) 4:325 (1842), defined
this structure as “ A small point (which) terminates the petioles 'whether
common or partial, in all or nearly all Mimoseae ” The present author
has modified Bentham’s usage of this term: as applied here the terminal seta
appendage which terminates the primary axis of a bipinnate leaf
(Bemharns “common petiole”). It does not refer to the apical portion
of the secondary axis of the leaf (Bentham’s “ partial petiole ”).
4. Areole: an area which occurs on each face of most Mimosoideae seeds
generally more or less elliptic or oblong and bounded by a fine line (the pleuro-
gtatn). (bee J. P. M. Brenan, Leguminosae-Mimosoideae in Flor Tron E
Africa, p. I (1959) tor a definition of areole, and E. J. H. Corner The Lesu-
gmmT I :l 17-150 (1951) for a definition of pleuL
Acacia megacephala B. R. Maslin sp. nov. (Figure I)
. ereclus diffusus 1-2 m alius, uni- vel niulti-caulis. ramulis .saepe pendulis modice
ad dense pilosis; cortex laevis, ad basin cinereo-brunneus. in ramulis rufo-brunneus’
axtilares. 5-15 mm longae, solitariae (saepe carentes), glabrae vel ad basin nilosae Srinulnf*
mmummmsrnm
breviter pilosum. BraclecAae dim^^^ eapilatum, glabrum vel
receplacidi dispositae mfltae obb gac ca’ I mm bne'.e acuta;: Tt '
(2) bracleolae superiores spiraliter disposhae erec aT ' ®
longi, Udine 4/5 ejus connatus tubo glabro vel {parsin piloso”?obis''ob^^^^ turbinatus.
c.liolalis; corolla 2-3 mm longa, lonfitudine 3/4 eiurconnata' /f "Pf*™
ovarium sessile, glabrum Leeumen 'ineare ad ui ® 12-granulana;
pagina parum undulata. bSr n^argintbus S ferl°rfT’ ^ s'abrum
tudinalia. oblonga 3^ x ..5-2.0 mm, fusca; funiculus p^erumque 2“ius.“u!“"®"
8 Aug.^mTO;^r/t‘' SS 6767hoL":%ER™!'i'o;"K,'°M^ Australia,
254
Figure l^Acacia met’acephala sp. nov. A— Portion of stem. B Portion of rachis showing
prominent midrib. C— Node with pinnae removed showing spine (sp), gland (g), terminal
seta (t), and stipule (s). D— Flower. E— Flower head. F- Recptacle (r) with di-
morphic bracteoles (b). G — Pod. H-— Young seed in pod. I - Seed showing tunicle
variability. . .
A-F from B.R. Maslin 676; G-H from B. R. Maslin 684; 1 Irom Phillips s.n.
Erect, dift'use, single-or multi-stemmed shnih, 1-2 m tall: branchlets
often pendulous, moderately to densely pilose; bark smooth, grey /brown at
base, red /brown on branches. Spines axillary, 5-15 mm long, solitary (often
absent from some nodes), glabrous, or pilose towards the base. Stipules
very narrowly triangular, 2-4 mm long, normally shortly pilose. Leaves
bipinnate; pinnae I pair; petiole very short, ca. 1 mm long; terminal seta very
255
narrowly triangular, 3-4(5) mm long, shortly pilose; gland arising at junction
of pinnae, stipitate, 4 equalling terminal seta, dilated at apex, glabrous or
shortly pilose; racliis 4-6 mm long, acute, sparsely pilose below, glabrous
and with a prominent midrib above; pinnules 4-6 pairs, narrowly obovate,
4-6 X 2-3 mm, obtuse, flat, inconspicuously 1 -nerved, smooth, glabrous,
dull green (at maturity). Flower heads large, globular, 8-10 mm diameter at
anthesis, with 80-90 flowers. Peduncles axillary, solitary or in pairs, greatly
exceeding the leaves, 1 5-25 mm long, glabrous. Receptacle capitate, glabrous
or shortly pilose. Bracteoles dimorphous; (1) lower series (arranged in a
single ring at base of receptacle) reflexed, oblong, ca. 1 mm long, acute, glabrous
except at apex; (2) upper series (spirally arranged) erect, ca. 1-5 mm long,
clavvs linear, normally glabrous, expanded into sparsely ciliolate, somewhat
concave, pear-shaped laminae. Flowers 5-merous; ealy.x about 2/3 length
of corolla, narrowly turbinate, connate for 4 5 its length, lube glabrous or
very sparsely pilose, lobes oblong obtuse sparsely ciliolate and somewhat
incurved at apex; corolla 2-3 mm long, connate for 3/4 its length, glabrous;
poUinia 12-grained; ovary sessile, glabrous. Legume linear to narrowly oblong,
25-50 X 4 mm. glabrous, surface slightly undulate, dark brown; margin thick-
ened, pale coloured, almost straight. Seeds longitudinal, oblong, 3-4 x I •5-2 0
mm, brown; funicle yellow, usually with 2 short folds, gradually expanded
into a thickened aril; areole ± oblong, 2 •5-3 0 x I 0 mm; pleurogram con-
tinuous.
Distribution: Western Australia; apparently confined to sandy or loamy soils
in an area extending from about 10 miles east of Geraldton eastwards to the
Greenough River. (Figure 4)
10 mi E of Geraldton. IV. E. Blackall 2755; 19 mi E of Geraldton, A. S. George 9220; 17 mi E
of Geraldton. J. iV. Green 474; 19 mi E of Geraldton on the road to Mullewa, B. R. Mastin
()84\ Northern Gully (E of Geraldton), G. Phillips .s.n.; ca. 50-5 mi W of Mullewa on the road
to Geraldton. M. D. Timlale 1328.
Flowering period: Augusl-September.
The presence of stipitate glands, axillary spines, and unijugate leaves
relates this species to A. lasiocarpa Benth. and more closely to A. pulehella
R.Br. However A. inegacephala is readily distinguished from both of these
species by its very long peduncles, prominently capitate receptacles, and its
large flower heads each bearing 80-90 narrowly turbinate flowers. Vegetatively
A. inegacephala is distinguished from A. pulehella by its one-spined nodes
(A. pulehella normally has 2 spines per node), longer narrower stipules and
terminal setae, and somewhat larger pinnules. The flat pinnules and longer
glands readily separate it from A. lasiocarpa.
Acacia echinata B. R. Maslin sp. nov. (Figure 2)
FriUe.x nanus, ramosissimus, compaclus, pulvinatus, 2-5 cm altus, 50 cm vel major diam.
partibus yegetativis sparsim striglllosis vel glabris; ramuli ± erect!, spinescentes, teretes;
spinae axillares nullae. Slipuiae anguste triangulares, 0-5-1 mm longae, praecipue ad basin
aliquantum incrassatae. Folia bipinnata; pinnae unijugatae; peiiohis prominens, ca. 2 mm
longus, teres, ad basin parum contractus; .wta terminalis 0-5-1 mm longa, aliquantum in-
crassata; glans in pagina supera petioli plerumque ad apicem, parva, sessilis, rotunda; rhachis
2-3 mm longa, teres, apice aliquantum complanato parum expanso, obtuso; pinnulae 3^-
jugae, anguste oblongae ad assymetricc anguste obovatae, 2-3 x 1 mm, obtusae, planae,
inconspicue 1-nervatae, laeves. Capitula florentia parva, globulosa, sub anthesi 3-4 mm’
diam., Horibus 12-15. Per/t/HCM// axillares, solitarii 10-15 mm longi, sparsim strigillosi ad
glabri. Reccpiaciilnm globulosum. glabrum. Bracleolae minus quam I mm longae, un-
guibus brevibus glabris, in laminas concavas obtusas ciliolatas expansae. Florae 5-merae;
ra/i-.v longitudine 1/2 corollae partes aequans, turbinatus, longiludine 3/4 ejus connatus
tubo glabro, lobis obtusis ciliolatis ad apices parum incurvatis; corolla longitudine I /2 ejus
connata, 1-5 mm longa. glabra; pollinia 12-granularia ; ovarium sessile parce papillatum.
Legumen anguste oblongum, ca. 10x3 mm. pagina undulata parce strigosa badia; marginibus
crassis, fere rectis, ferrugineis. Semina matura non visa.
Type: 6 miles east of Kukerin, Western Australia, 20 Dec. 1964 K Newbev 1620 (holo-
PERTH, iso: K. MEI,, NSW). '
256
Figure 2— Acacia echinata sp. nov. A— Portion of branch system. B— Node showing
petiole (p), stipules (s), terminal seta (t), and gland (g). C — Flower. D Flower head.
E— Receptacle (r) with bracteoles (b). F— Pod valves, side and top view. All from K.
Newbey 1620.
Dwarf, much branched, compact, cushion-like shrub, 2-5 cm high and
50 cm or more in diameter, vegetative parts sparsely strigillose to glabrous;
branchlets - erect, spinescent, terete; axillary spines absent. 5///7u/c^ narrowly
triangular, 0-5-1 mm long, somewhat thickened especially near the base.
Leaves bipinnate; pinnae I pair; petiole prominent, 3-4 mm long, terete,
slightly contracted at base; terminal seta 0-5-1 mm long, somewhat thickened;
vlaiul on upper surface of petiole (normally near the apex), small, sessile,
circular; rachis 2-3 mm long, terete, apex somewhat flattened and slightly
expanded, obtuse; pinnules 3-4 pairs, narrowly oblong to asymetrmally nar-
rowly obovate, 2-3 x ca. 1 mm, obtuse, flat, inconspicuously l-nerved, smooth.
Flower beads small, globular, 3-4 mm diameter at anthesis, with 12- 15 flowers.
Peduncles axillary, solitary, 10-15 mm long, sparsely strigillose to glabrous.
Receptacle globular, glabrous. Bracteoles less than 1 mm long, claws short,
glabrous, expanded into concave, obtuse, ciliolate laminae. Fbwetj 5-merous;
cahx about 1 /2 length of corolla, turbinate, connate for ca. 3 /4 its length,
tube glabrous, lobes obtuse ciliolate and slightly incurved at &pex- corolla
1 -5 mm long, connate for 1 /2 its length, glabrous; pollinia 12-gramed; orarr
sessile, very sparsely papillate. Legume narrowly oblong, ca. 10 x 3 mm,
with a dark brown undulate moderately strigose surface; margins thickened,
light brown, almost straight. Seed not seen in mature state.
Distribution: South-west Western Australia; known only from the type locality
which is a gravelly hill between Kukerin and Tarm Rock. (Figure 4)
6 mi W of Tarin Rock, F. LuUfitz 5901 ; 198 mile peg Dumbleyung-Lake Grace road, A. S
George 5800.
Flowering period: December-January.
The cushion-like habit, the short spinescent branchlets, and the single
pair of pinnae borne on a long petiole make A. echinata a very distinctive
species within the series Pulchellae.
257
Figure 3 — Acacia moirii E. Pritzel subsp. recurvistipida subsp. nov. A — Portion of branch.
B — Leaf with gland (g), and terminal seta (t). C — Flower. D — Flower head. E — Recept-
acle with bracteoles. F — Pod valve.
A-E from A. S. George 6349; F from B. R. Maslin 525.
Acacia moirii E. Pritzel subsp. recurvistipula B. R. Maslin subsp. nov. (Figure 3)
A subspecie moirii stipulis spinescentibus recurvis; rhachidi 5 mm longa apice recto;
pinnulis 3-5-jugis, parum incrassatis, enerviis vel inconspicue l-nervatis, laevibus vel sparsim
puberulis aliquantum glaucis, differt.
Type: 9 miles north of ‘ The Humps ' (which is ca. 10 miles due north-east of Hyden), Western
Australia, 15 July 1970, B. R. Maslin 570 (holo: PERTH).
Differs from A. moirii E. Pritzel subsp. moirii chiefly in vegetative charac-
ters: stipules recurved, spinescent; rachis 5 mm long, apex straight; pinnules
3-5 pairs, scarcely thickened, nerveless or obscurely 1-nerved, glabrous to
sparsely puberulous, somewhat glaucous.
Dwarf, erect, compact shrub, 20-40 cm tall; branches ± flexuose, an-
trorsely puberulous, axillary spines absent. Stipules spiny, 3-6 mm long,
recurved, sparsely antrorsely puberulous, persistent. Leaves bipinnate;
pinnae 1 pair; sparsely antrorsely puberulous to glabrous; petiole very short,
ca. 1 mm long; terminal seta spiny, 2-3 mm long, straight or somewhat re-
curved, persistent; gland on upper surface of petiole, circular, sessile; rachis
ca. 5 mm long, terete, apex straight or slightly recurved -f expanded; pinnules
3-5 pairs, narrowly oblong to obliquely elliptic, 2-3 x ca. 1 mm, obtuse flat
nerveless or inconspicuously 1-nerved, smooth, slightly thickened. Flower
heads globular, ca. 5 mm diameter at anthesis, with ca. 15 flowers. Peduncles
axillary, solitary, 5-10 mm, mostly retrorsely puberulous. Receptacle short
ovoid, ca. I mm long, densely puberulous. Bracteoles ca. 1 mm long claws
glabrescent, gradually expanded into indexed, pilose laminae. Flowers
258
5-merous; calyx ca. 1 /2 length of corolla, turbinate, connate for 3 /4 its length,
5-nerved, tube glabrous, lobes oblong obtuse ciliolate (often invested with a
few additional long straight hairs) and somewhat incurved at apex; corolla
1-5-2 mm long, connate for ca. 1 /2 its length, 5-nervcd, usually glabrous
(sometimes invested with a few long straight hairs at apex ); poUinia 12-grained;
ovary sessile, sparsely papillate. Legume — see below. Seed n.v.
259
Distribution: South-west Western Australia; from Kukerin to Hyden and
eastwards to about Lake King. (Figure 4)
8 mi N of ‘ The Humps B. R. Maslin 569; ca. 4 mi S of Kulin, B. R. Mastin 525 ; I97J mile-
peg, E. of Dumbleyung, A. S. George 6349; 16 km W of Lake King township, P. G. Wilson
7159; 37 km E of Lake King, D. Young 129.
Flowering period: May-July.
A. moirii subsp. recurvistipida differs from the type subspecies chiefly in
vegetative characters. Although the stipules of subsp. moirii are prominent,
and sometimes thickened at the base, they are never spiny or recurved as in
the new subspecies. In addition, the rachis of subsp. moirii is about twice as
long as in subsp. recurvistipida and has a more prominently recurved tip.
The pinnules of the two subspecies are quite different; in the type subspecies
they are thicker, greater in number, and have a prominently raised midrib
(a character never found in subsp. recurvistipida), they are generally less glau-
cous, and they have a denser indumentum.
This subspecies usually occurs in open places on sandplains where it
grows as a small erect compact shrub. However, on a few occasions it has
been found growing among litter beneath tall shrubs such as Casuarina pinaster
C. A. Gardner. In these latter cases subsp. recurvistipida assumes a semi-
prostrate and somewhat openly branched habit; in addition its pinnules and
peduncles are somewhat enlarged, while the stipules and terminal seta are not
so rigid.
No seed of subsp. recurvistipida has been collected. The following is a
description of a few dehisced fruiting valves caught in the branches of B. R.
Maslin 525: narrowly oblong, 25 x 5 mm, curved, densely pilose, dark brown
slightly undulate, margins thickened.
The distributions of the two subspecies do not appear to overlap. A.
moirii subsp. moirii has the more southerly range, extending from about Jerra-
mungup to Ravensthorpe while subsp. recurvistipida occurs further inland.
260
New species and subspecies of Casuarina in Western Australia
By L. A. S. Johnson*
Abstract
The following taxa are described and discussed: C. sclerodada sp. nov.,
C. campestris Diels subsp. lessellata (C. A. Gardn.l comb, et stat. nov., C.
campeslris subsp. eriochlamys subsp. nov., C. campestris subsp. grossa subsp.
nov. and C. crislala Miq. subsp. pauper (F. Muell. ex Miq.) comb, et stat. nov.
Introduction
The following descriptions of new taxa, together with the reduction of a
supposed species to subspecific rank, are published so that the names will be
available for use in the forthcoming treatment of Casuarinaceae in “ The
Flora of Western Australia It is intended to deal with the general taxonomy
of the family elsewhere.
For morphological reasons certain terms are employed here and elsewhere
in my Casuarina studies, for which those more traditionally used in Casuarina
are indicated in parenthesis: leaf lamina (“tooth", "scale-leaf”), article
(“ internode ”), phyllichnium (“ ridge "), infructescence (“ cone ”).
Casuarina sclerodada L. Johnson, sp. nov.
[Greek: skleros hard, klados = branch]
Friitex dioicus 1-3 m altus, saepe trunco singulo. ramis saepe arcuatis, ramulis pendenti-
bus. Rami persistentes novelli ramulis deeiduis aliquanto similes sed internodiis brevioribus
distinct! vetustiores mox lignosi cortice plus minusve sulcato. Ramiili decidui multiarticulati,
penduli sed duri: verticilli plerumque (9-)10-l 1-lbliati; laminae foliariim C dentes ”) primo
circiter I -0-1 -5 mm longae, demum (fissione vaginarum) usque ad 2-0 mm longae, elongato-
triangulares, erectae, brunneae nigrescenlesquc, marginibus praeserlim basin versus plus
minusve lanuginoso-ciliatis; arliciili (" inlernodia ) (1—) 2— 4(-5) cm longi, plerumque O ' 7-0 9
mm crassi, in* sicco cinerascenlcs: phrllichnia subconvexa (in sicco), baud carinata, pilis sul-
corum superficie manifestis sed non protrudentibus. Infiorescenliae masciilae in ramis per-
sistentibus se.ssiles (saepe cum ramulis vegetativis in verticillis isdem), O'3-O'l (vel plus) cm
longae- verticilli bractearum crebri, ad 1-2 mm longi (laminis inclusis); laminae bractearum
illis foliorum similes sed breviores, circiter 0-5 mm longae; bracteolae persistentes, stipitatae,
elongato-cvmbiformes, acutae, apicibus ad anlhesin plus minusve^ expositis, marginibus
versus apicem minute ciliolatis; tepalum I (abaxiale deticiens), adaxiale oblanceolatum, vix
cucullatum, antheram partim includens; filamsntum I -5 mm attingens; anlhera rubens, api-
culata circiter 0-7 mm longa. Infiorescenliae foemineae in ramis persi.stentibus sessiles,
ovoideae, 6-8 mm longae, bracteis tomentosis. infructescentme ("strobih ’) sessiles, sub-
globosae vel crasse ellipsoideae, apice truncate, I 5-3-0 cm longae, 1 -5-2-5 cm diametro.
18-22-stichae verticillis fertilibus plerumque 8-12; bracteae tenues apice tnangulari tomen-
toso minute a'piculato exposito; bracteolae (" valvae ") maturitate lignosae, apice cras_so con-
vexoque obtuso, plus minusve glabrato et sine protuberatione separata. Nux tota 5-8 mm
lonca corpore mature atro vix nitenti, ala expansa quam corpora saepe aliquanto longiore,
translucent! basi styli excepta, ad apicem suboblique rotundata apiculataque. Chromoso-
mata somatica 50-52.
Hololvpiis c. 40 km SSE. of Caiguna (c. 3-2 km in from sea), W.A. 2-3 m, very
arching and drooping, no 3 seen, on limestone plain with mallee and scrub, I,ix.l967, L. A. b.
Johnson 2155, NSW 132432, isotypo in PERTH ponendo.
Dioecious shrub 1-3 m, often with a single trunk, the branches often arching
and the branchlets drooping. Young permanent branches rather similar to
the deciduous branchlets but distinguishable by their shorter articles (
nodes ”), the older ones soon becoming woody with more or less furrowed.
* National Herbarium of New South Wales, Royal Botanic Gardens, Sydney, N.S.W.
2(X)0.
261
hard, grey bark. Deciduous (“ foliage ”) branchlets many-jointed, pendulous
but hard; whorls usually (9-) 1 0- 1 1 -leaved (“ -toothed ”); leaf laminae (“ teeth ”)
at first c. 1 0-1 -5 mm long but finally apparently up to 2 0 mm long (by splitting
of the sheaths), elongate-triangular, erect (appressed to succeeding article),
brown and becoming blackish, margins more or less woolly-ciliate especially
towards the base; articles (“ internodes ”) (l-)2-4(-5) cm long, usually 0-7-0'9
mm thick, becoming ashy-grey when dry; phyllichnia (“ridges ) slightly
conve.x (when dry) but not keeled, the hairs of the furrows visible on the surface
but not protruding. Male inflorescences sessile on the permanent branches
(often with vegetative branchlets in the same whorl). 0-3-1 0 (or more) cm
long; whorls of bracts crowded, to 1-2 mm long (including the laminae);
laminae of the bracts (“ teeth ’’), similar to those of the leaves but shorter,
about 0-5 mm long; bracteoles persistent, stipitate, elongate-cymbiform. acute,
their apices more or less e,\posed at anthesis, the edges minutely ciliolate
towards the apex; tepal I (the abaxial one lacking), adaxial, oblanceolate,
scarcely hooded, partly enwrapping the anther; filament attaining 1-5 mm;
anther reddish, apiculate, c. 0 • 7 mm long. Female inflorescences sessile on the
permanent branches, ovoid, 6-8 mm long, with tomentose bracts. Infructes-
cences (“ cones ”) sessile, subglobose or thickly ellipsoid, truncate at the apex,
1-5-3-0 cm long, 1 •5-2-5 cm in diameter, 18-22-stichous {i.e. with alternate
9-1 1-membered whorls), fertile whorls usually 8-12; bracts thin with the tom-
entose minutely apiculate apex exposed at the cone surface; bracteoles
(“ valves ”) woody at maturity, the apex thick and convex, obtuse and without
a separate dorsal protuberance. Nut 5-8 mm long overall, the body black and
scarcely shining when ripe, the wing expanded and usually somewhat longer
than the body, translucent (but brownish) apart from the style-base, sub-
obliquely rounded and apiculate at the apex. Chromosome No. 2n =- 50-52.
Distribution: Western Australia, from the Borden district to the western part
of the Great Australian Bight.
Specimens examined: Western Australia: 312 miles (499 km) on the Lake King-Ravensthorpe
road, E. M. Bennett 3120, 16. i. 1970 (S and $, apparently separate plants) (PERTH, NSW);
17 miles (27 km) E. of Ravensthorpe on Esperance road, B. G. Brings 442, 443, 444a, ll.ix.l966
(NSW); Mt Ragged, near SW. base, A. S. George 2110, 7.xii.l960 (PERTH); Salmon Gums
C. A. Gardner, v.1924 (PERTH); Great Bight, Carey (MEL); 6 miles S. of Toompup, Newbey
1249 (PERTH); and the Holotype. Further collections from the Caiguna area by R. F.
Parsons seen, but details not recorded (AD).
This sharply defined species is probably nearest to C. acutivalvis F. Muell.,
from which it differs in the arching habit of growth (more pronounced in
female plants, A. S. George pers. comm.), short sessile male inflorescences,
the smaller and always sessile infructescences with obtuse bracteoles, the coarser
branchlets and other vegetative parts, as well as in numerous fine details. It
differs clearly also from other species of this group, such as C. campestris
Diels and C. dielsiana C. A. Gardn. in habit, male inflorescences, infructescences,
and branchlets.
C. scleroclada has been little collected but is common enough in places,
for instance in the type locality towards the Bight coast from Caiguna. It was
first recognized by the author some twenty years ago from the Gardner and
the rather fragmentary Carey collections but for many years it was impossible
to gain any field knowledge of it or to obtain suitable material for chromosome
number determination.
Unlike the diploid C. acutivalvis, C. campestris, and C. helmsii Ewart
& Gordon, all of which have In 24 (Barlow 1959), and C. dielsiana (2n - 28),
C. scleroclada (at least from the type locality) is at the tetraploid level with
2n = 50-52.
262
Casuarina campestris Diels in Bot. Jahrb. 35:126 (1905)
This widespread species is fairly uniform over much of its range but in
the Mt. Singleton and Kalgoorlie districts, as well as towards the southern coast,
populations occur which appear to be best retained within C. campestris but
differ sufficiently to warrant subspecific rank. C. cUelsiana C. A. Gardn. also
greatly resembles C. campestris in vegetative features but differs in the infruc-
tescences, the peculiar short, more or less globose male inflorescences, and in
chromosome number (Barlow 1959).
Thus C. campestris can be regarded as comprising tour subspecies which
show some morphological intergradation and some geographic overlap. For
some purposes, and with intermediate material, it may be convenient to refer
simply to C. campestris, but the following key indicates the more characteristic
features of the four races. Specimens without infructescences may be difficult
to determine unless they are associated with fruiting material from the same
population.
Key to Subspecies
1. Surface of fruiting bracteoles divided into 4 “ protuberances ” (Infructescences pedunculate,
glabrescent, bracteoles protruding, phyllichnia markedly convex when dry, leaf-whorls
mostly 9-nierous) !• subsp, tessellata
1*. Surface of fruiting bracteoles not divided.
2 Infructescences mostly sessile or subsessile (fruiting peduncles mostly 0-4 mm), mostly
1 -3^-o X I -2-1 -hem, glabrescent when ripe. Bracteoles protruding, phyllichnia mark-
edly convex when dry (leaf-whorls 7-9-merous) 2. subsp. campestris
2* Infructescences on peduncles of 4-15 mm, either greyish-pubescent to tomentose when
ripe or mostly I •8-2-8 cm in diameter. Bracteoles flush or protruding. Phyllichnia
only slightly convex when dry
3 Infructescences mostly 2-0-3-5 x I -3-1 -6 cm, with fruiting bracteoles L flush with
general surface and tawny to greyish pubescent or tomentose. Articles 5-12 mm long,
0-8-0'9 mm diam. (when dry), leaf-whorls 8-9-merous 3. subsp. eriochlamys
3* Infructescences mostly 2-0-5-2 x 1 ■8-2-8 cm, with fruiting bracteoles markedly
protruding, tawny pubescent at first but glabrescent when fully ripe. Articles 8-15
mm long, 0-9-1 -I mm diam. (when dry), leaf-whorls 8-11-merous 4. subsp. grossa
1. subsp. campestris
Lectotvpus: Watheroo, L. Diels 2038, 31.xii.l900 (B). This is one of four
syntypes and is a specimen with mature infructescences which are characteris-
tically sessile, slender, and glabrescent. The Casuarinaceae in the Berlin
Herbarium fortunately survived the 1939-1945 war.
This is the very wide-ranging and familiar form of C. campestris found
throughout the wheat belt of Western Australia on sandplains and laterite.
Occasional individuals may be found with over-sized infructescences, and there
are forms near the southern coast which may approach subsp. grossa in this
respect, though the infructescences are usually sessile or subsessile. Inter-
grading to subsp. eriochlamys appears to occur in the Coolgardie district.
There is geographic overlap with subsp. grossa in the Norseman-Esperance
area (see below).
2. subsp. tessellata (C. A. Gardn.) L. Johnson, comb. et. stat. nov.
C tessellata C A Gardn. in J. Roy. Soc. W. Austral. 22:119 (1936). Holo-
typiis: Summit of Mt. Singleton. C. A. Gardner 2217, 9.vii.l931. “erect,
10-15 ft. high, red loamy soil ”, ■? with cones (PERTH, isotype in K).
1 have seen a number of collections, but all are from the type locality and
they include no male material. A possibility remains that male inflorescences or
chromosome number may support its retention as a species, but on the known
characters it appears to differ no more from C. campestris subsp. campestris
263
than do the other subspecies here recognized. Subsp. tesseUala is distinguished
by the combination of characters listed in the Key. The peduncles range from
7 to 15 mm in length and the infructescences are similar in shape and size
(but not bracteoles) to those of subsp. campestris, which subsp. tessellata also
resembles in branchlet dimension and surface.
3. subsp. eriochlamys L. Johnson, subsp. nov.
[Greek: erion ^ wool, chlamys = mantle]
Afliai/i ramuloruni 5-12 mm long!, 0-8-0 -9 mm crassi, phyllichniis subconyexis, verti-
cillis plerumque 8-9 folialis. hiflorescenliae masculae eis subsp. campestris similes. //;-
fructescentiae (“ strobili ’') in pedunculis 7-13 mm longis.maturae circiter 2 0-3 -0 cm longae,
1- 2-1 -6 cm diametro; bractearum facies externa ea subspecierum alterarum crassior; brac-
teolae (" valvae ") baud protrudentes, ad maturitatem pubescentes vel cano-tomentosae.
Hohtypus: NSW 61832, Comet Vale, J. T. Jursoii 255, viii.1917, ? with inflorescences and
infructescences.
Branchlet articles 5-12 mm long, 0 -8-0 -9 mm thick, phyllichnia (“ ridges ”)
somewhat convex but not prominently so, whorls with 8-9 leaves (“ teeth ”).
Male inflorescences like those of subsp. campestris. Infructescences (“ cones ”)
on peduncles of 7-13 mm, when mature 2 -0-3 -0 cm long, 1 -3-1 -6 cm in dia-
meter; the outer face of the bracts thicker than in the other subspecies: brac-
teoles (“ valves ■’) not protruding (flush with the general cone surface), tawny
to greyish or whitish pubescent or tomentose at maturity.
This subspecies is represented by a number of collections from Comet Vale
(of which Royce 4444 and Gardner 7969 in PERTH are good examples). Some
specimens of subsp. campestris from the Coolgardie district show some approach
to subsp. eriochlamys in having a denser and more persistent indumentum on
the infructescences and less convex phyllichnia than is usual in the type sub-
species. Again, subsp. grossa shares certain characters (pedunculate infruc-
tescences, non-prominent phyllichnia) with subsp. eriochlamys and in the north-
ern part of its range may approach the latter subspecies.
4. subsp. grossa L. Johnson, subsp. nov.
[Latin: grossus = coarse, thick]
Articiiii ramulorum 8-15 mm longi, 0-9-1 - I mm crassi, phyllichniis planis vel subcon-
vexis, verticillis 8-1 l-foliatis. Infloresccntiae masculae eis subsp campestris similes sed
aliquanto crassiores. Infructescentiae (“strobili”) in pedunculis 4-15 mm longis, maturae
2 - 0-5 -2 cm longae, 1 -8-2-8 cm diametro; bracteolae (“ valvae ") valde protrudentes, primum
fulvo-pubescentes sed ad maturitatem glabrescsntes.
Hohtypus: NSW 58480, 51 miles (9 km) N. of Norseman, L. A. S. Johnson W177, 18.xii.l960,
" bushy shrubs to 2-2-5 m around granite tors with Eucalyptus websterana. Uniform popu-
lation. Dioecious here. C. heimsii nearby ”. Isotype in PERTH.
Branchlet articles 8-15 mm long, 0-9-1 - 1 mm thick, phyllichnia (“ ridges ”)
flat or somewhat convex but not prominently so, whorls with 8-1 1 leaves
(“ teeth ”). Male inflorescences like those of subsp. campestris but somewhat
thicker. Infructescences (“cones") on peduncles of 4-15 mm, when mature
2-0-5-2 cm long, 1 -8-2-8 cm in diameter; bracteoles (“valves”) strongly
protruding, at first tawny-pubescent but glabrescent at maturity.
This subspecies grows characteristically around granite tors in the Norse-
man district, a habitat rather different from those of subsp. campestris. There
are a number of collections from this area and habitat but there are also forms
with large infructescences and coarse branchlets in various other south-eastern
localities which seem to connect subsp. grossa with subsp. campestris.
More extensive collecting and exploration may make it clearer whether
these four races are in fact best treated as subspecies.
264
Casuarina cristata Miq., Rev. Crit. Cas. 70, t.lO (1848).
This widespread species will be dealt with elsewhere but it is convenient
at this stage to establish formally the two subspecies into which it may be sub-
divided.
1. subsp. cristata
This comprises the populations in Queensland (except for a small area of
subsp. pauper in the far south-west, isolated from other Queensland occurrences
of the species) and in New South Wales cast of a line from west of Moree to
Lake Cargelligo and Barmedman. In this area, where it is known as Belah ”,
it occurs on heavy grey or black soils with calcium carbonate in the lower
horizons. The type locality is at “ Fields Plains ”, now the Condobolin dis-
trict. For some 50 km or so to the west of this line there are occasional occur-
ences of forms intermediate between subsp. cristata and subsp. pauper.
2. subsp. pauper (F. Muell. ex Miq.) L. Johnson, comb, et stat. nov.
C. pauper F. Muell. ex Miq. in Ned. Kruidk. Arch. 4:98 (1859).
This subspecies, the ” Black Oak ” of South Australia, extends from western
New South Wales (west of the transitional zone to subsp. cristata) through
north-western Victoria and South Australia (the type locality is in Eyre Pen-
insula) to southern inland districts of Western Australia, where it appears
sometimes to hybridize with C. obesa Miq.
C. cristata subsp. pauper also occurs on soils with a calcareous subsoil
but these usually have a lighter-textured and more reddish or brownish surface
horizon than those on which subsp. cristata occurs. It differs from subsp.
cristata in the usually thicker and more waxy-surfaced branchlets and the
shorter fruiting bracteoles of the infructescences, which are also often more
persistently tawny-pubescent. The trees are usually smaller and of poorer
form.
C. obesa is more closely related to the eastern Australian C. glauca Sieber
ex Spreng. and, like that species, occurs on more or less saline soils. It is
common in Western Australia but in South Australia and New South Wales
it is known to persist in only two localities, whilst in Victoria it is represented
only by one old unlocalized collection.
I am grateful to the past and present officers in charge of the herbaria
from which specimens have been examined and to Dr R. F. Parsons for the
opportunity of e.xamining his collections. My colleague Dr Barbara Briggs
has kindly made available the result of her investigation of the chromosome
number of C. scleroclada. Mrs Eleanor Bennett has been most helpful in
the provision of material and information.
Reference
Barlow, B. A. { 1959)— -Chromosome numbers in the Casuarinaceae. Austral. J. Bot. 7:230-
237.
265
New taxa and new combinations in Western Australian
Pittosporaceae
By Eleanor M. Bennett
Abstract
It is proposed that the genera Marianthus and BiUardiera be combined
under BiUardiera. The necessary new combinations are made, and three new
varieties described. Two new varieties are described in the genus Cheiranthera.
A new combination, using the earliest available epithet, is provided for the
solitary species of Pronaya.
Introduction
The genera Marianlhus and BiUardiera are here combined under Billard-
iera. These genera have been distinguished by former workers on their fruit
which in BiUardiera is a berry and in Marianthus is a capsule. This division
does not appear to be a natural one since a separation based on floral characters
would provide a quite different arrangement of species. The use of fruit
characters for delimiting these two genera also makes impossible the ready
identification of flowering material, as an examination of the ovary provides
no clue to its future form and in fact several species have been, and continue
to be, incorrectly placed.
In addition two new varieties of Cheiranthera are described, one in each
of the two species which occur in Western Australia.
In Pronaya, the name P. elegans Hueg., which has been in common use,
is antedated by the name Spiranthera fraseri Hook, and a new combination
is made accordingly.
Species have been accepted after consulting type specimens, descriptions
and dates of publication, occasionally resulting in the changing of previously
accepted names.
These new combinations and varieties are being published prior to a
revision of the Western Australian species of Pittosporaceae in a forthcoming
Flora part.
BiUardiera
BiUardiera Sm., Spec. Bot. Nov. Holl. l:t.l (1793). Type: B. scandens Sm.
Lahillardiera Roem. et Schult., Syst. Veg. 5:330 (1819). nom. illeg.
based on above.
Marianthus Hueg. ex Endl., Enum. PI. Hueg. 8 (1837). Type: M. Candidas
Hueg. ex Endl.
Oncosporum Putterl. in Endl. et Fenzl., Nov. Stirp. Dec. 9 (1839).
Type: C. bicolor Putterl.
Calopetalon J. Drumm. ex Harv., Hooker's Journ. Bot. Kew Card. Misc-
7:52 (1855). Type: C. ringens J. Drumm. ex Harv.
Cyathomiscus Turcz., Bull. Soc. Imp. Nat. Mosc. 36/1:562 (1863).
Type: C. cuneatum Turcz.
BiUardiera laxiflora (Benth.) E. M. Bennett comb. nov.
Marianthus ? laxiflorus Benth., FI. Austral. 1:119 (1863). Lectotype:
Herb. Mueller, Porongurup Range, W. Australia, (holo:MEL).
BiUardiera gracilis Diels, Bot. Jahrb. 35:213 (1904). Type: King George
Sound, Diels 2259 and Pritzel 252 (syn:PERTH).
266
Billardiera villosa (Turcz.) E. M. Bennett comb. nov.
Oncosporum villosum Turcz., Bull. Soc. Imp. Nat. Mosc. 27:365 (1854).
Type: Western Australia, Drummond 5th Coll., n.242 (iso.MEL).
Oncosporum mkrophyllum Turcz., loc. cit. Type: not given.
Marianthus mkrophyllum (Turcz.) Benth., El. Austral. 1:117 (1863).
M. rhytidosporus F. Muell., Fragm. 2:145 (1861). Type: Near Esperance
Bay, W. Australia, A/a.vM’c// (holo:MEL).
Billardiera granulata (Turcz.) E. M. Bennett comb. nov.
Oncosporum graniilaium Turcz., Bull. Soc. Imp. Nat. Mosc. 27:366 (1854).
Type: Western Australia, Drummond 3rd Coll., n.210 (iso:W).
Marianthus granulalus (Turcz.) Benth., El. Austral. 1:118 (1863).
M. parviflorus F. Muell., Fragm. 2:144 (1861). Type: Porongurup Range,
W. Australia, Maxwell (holo:MEL).
Billardiera Candida (Hueg. ex Endl.) E. M. Bennett comb. nov.
Marianthus Candidas Hueg. ex Endl., Enum. PI. Hueg. 8 (1837).
Type: Western Australia, Huegel (holo:W).
Billardiera coeruleo-punctata (Klotsch) E. M. Bennett comb. nov.
Marianthus coeruleo-punctatus Klotsch in Link, Klotsch et Otto, Ic.
PI. Rar. 28 t. 12 (1840). Lectotype: Western Australia, 1839, Drummond
(iso :W).
Pronaya pedunculosa Turcz., Bull. Soc. Nat. Mosc. 36:560 (1863).
Type: Western Australia, 1839, J. Drummond (\\o\o:'^).
Billardiera drummondiana (Putterl.) E. M. Bennett comb. nov.
Oncosporum drummondianum Putterl. in Lehm., PI. Preiss. 1:194 (1844).
Lectotype: Western Australia, Preiss 1288 (holo:W).
Marianthus drummondianus (Putterl.) Benth., FI. Austral. 1:119 (1863).
Billardiera dorrienii Domin. Mem.Soc.Sc.Boheme 2:24 (1923) (holo:K,
photo seen).
B. drummondiana var. collina E. M. Bennett var. nov. (L. occurring
on hills)
Folia, caules, calyces et pedicelli dense hirsuti pilis longis unicellulosis etiam
breves multicellulosis. Pedicelli 4-10 (15) mm longi.
Type: Bushmead-Kalamunda area, W. Australia, 19 Sep. 1966, F. W. Hum-
phreys 24, (holo:PERTH).
Leaves stems, calyces and pedicels densely hirsute with long unicellular
and short multicellular hairs. Pedicels 4-10(15) mm long.
The typical variety differs from var. coUina in having only unicellular
hairs scattered on the leaves, stems and calyces and in having long pedicels.
It is found around Cranbrook and Manjimup while var. collina is found in the
Darling Ranges.
Billardiera bicolor (Putterl.) E. M. Bennett comb nov.
Oncosporum hicolor Putterl. in Endl. et EenzL, Nov. Stirp. Dec. 9 (July
1839). Type: Western Australia, Huegel (n.v.).
Marianthus pictus Lindl., Sketch Veg. Swan Riv. Col. 22 (Dec. 1839).
Type: Western Australia (holo:CGE, photo seen).
B. bicolor var. lineata (F. Muell.) E. M. Bennett comb, et stat. nov.
M. lineatus F. Muell., Fragm. 1:27 (1859). Type: Murchison River,
W. Australia, F. Mueller. (holo:MEL; iso:W).
In var. lineata, the pedicel is stout, the corymbs are shorter than or just
exceed the leaves and the capsule is cartilaginous. The typical variety has a
slender pedicel, corymbs which exceed the leaves and a crustaceous capsule.
Billardiera erubescens (Putterl.) E. M. Bennett comb. nov.
Maricmthus erubescens Putterl. in Endl. et FenzI., Nov. Stirp. Dec. 60
(1839). Type: Western Australia, 1837, Roe {holo;W).
M . purpiireus Turcz., Bull. Soc. Nat. Mosc. 27:364 (1854). Type: Western
Australia, J. Drummond 4th Coll., n.96, (n.v.).
Billardiera ringens (Drumm. ex Harv.) E. M. Bennett comb. nov.
Calopetalon ringens Drumm. ex Harv., Hooker’s Journ. Bot. Kew Garden.
Misc. 7:52 (1855). Type: Chapman, W. Australia, J. Drummond (n.v.).
Marianthus ringens (Drumm. ex Harv.) F. Muell., Fragm. 1:218 (1859).
Billardiera sericea (Turcz.) E. M. Bennett comb. nov.
Pronaya sericea Turcz., Bull. Soc. Imp. Nat. Mosc. 27:363 (1854).
Type: Western Australia, J. Drummond 4th Coll., n.97 (iso:MEL).
Billardiera parviflora DC., Prodr. 1:346 (1824).
Type: Western Australia, Leschenault (holo:G-DC).
Marianthus tenuis Benth., FI. Austral. 1:119 (1863). Syntypes: Flinders
Bay, W. Australia, Collie (K, MEL); Geographe Bay, Leschenault (n.v.);
Cape Naturaliste, Oldfield
M. gracilis Ostenf., Dansk. Vidensk, Selsk. Biol. Medd. 312;67 PI. 9.
(1921). r>’/7e.- Yallingup Cave, 26 Sept. 1914, Oi/cw/c/r/ 1018. (n.v.)
B. parviflora var. guttata E. M. Bennett var. nov. (L. guttatus — spotted)
Petala caesia, fauce atro-cyaneo-maculata.
Type: Armadale-Jarrahdale, W. Australia, 3 Aug. 1960, F. C. J. LuHfitz
(holo;PERTH).
The var. guttata differs from the typical variety in the petals having dark
blue spots in the throat. It occurs in the southern areas of the Darling Range
whereas the typical variety is found near Augusta.
Cheiranthera
Cherianthera preissiana Putterl. in Lehm., PI. Preiss. 1;201 (1845).
Type: York, Preiss 1291 (iso; MEL).
C. parviflora Benth., FI. Austral. 1;I28 (1863). Type: Western Australia,
J. Drummond n.34 and 80 (syn;MEL).
C. preissiana var. planifolia E. M. Bennett var. nov. (L. planus, folium — flat
leaved)
Folia linearia, plana, glabra vel pilis sparsis. Surculi et folia juvenes pilis
sparsis.
Type: Nornalup, W. Australia, E. M. Bennett 3184, 19 Jan. 1970 (holo-
PERTH).
Leaves linear, flat, glabrous or with scattered hairs. Young shoots and
leaves with scattered hairs.
The typical variety differs from var. planifolia in having ovate, pubescent
leaves.
Cheiranthera filifolia Turcz., Bull. Soc. Nat. Mosc. 27;364 (1854).
Type: Western Australia, Drummond 4ih Coll, n.94, (iso;MEL).
C. tortilis F. Muell., Fragm. 2;79 (1860). Type: South-west of Western
Australia, F. Mueller (n.v.).
268
C. filifolia var. brevifolia (F. Muell.) Benth., FI. Austral. 1:128 (1863).
C. brevifolia F. Muell., Fragm. 1:97 (1858). Type: Phillips Range,
F. A/we//e>r(holo:MEL).
C. filifolia var. simplicifolia E. M. Bennett var. nov. (L. simplex, folium —
simple leaved)
Folia plana, linearia, acuminata.
Type: Pindar, W. Australia, C. A. Gardner 7766 (holo:PERTH).
Leaves flat, linear, acuminate.
This variety dilTers from the typical variety and from var. brevifolia (F.
Muell.) Benth. in having flat leaves. The other two varieties have terete,
canaliculate leaves.
Pronaya
Pronaya fraseri (Hook.) E. M. Bennett comb. nov.
Spiranthera fraseri Hook., Bot. Mag. 63, sub tab. 3253 (1836).
Type: Western Australia, Fraser (n.v.).
Campylantliera fraseri (Hook.) Hook., leones PI. 1: tab. 82 (Mar. 1837).
Pronaya speciosa Endl., in Endl. et ah, Enum. PI. Huegel: 9 (April 1837).
Type: King George Sound, W. Australia, Ferd. Bauer (holo: W).
Pronaya elegans Hueg., Bot. Archiv. tab. 6 (April-May 1837).
Tvpe: Western Australia, F. Bauer (n.v.). Billardiera elegans (Hueg.)
F. Muell.. PI. Indig. Col. Viet. 1 : 78 (1862).
P. fraseri var. minor (Benth.) E. M. Bennett comb. nov.
Pronaya elegans Hueg. var. minor Benth., FI. Austral. 1 : 126 (1863).
Type: South coast (W. Australia), R. Brown (n.v.).
269
A revision of the genus Lamarchea Gaudichaud
(Myrtaceae: Leptospermoideae)
By A. S. George
Abstract
The Australian genus Lamarchea Gaud,, containing two species and one
variety, is described and discussed. One new species is described, L. sulcata
sp. nov. from the west-central Australian deserts, and one new variety, L. hake-
ifolia Gaud. var. hrevijolia var. nov. from near Shark Bay, Western Australia.
Introduction
A revision of this small genus, previously thought to be monotypic, was
prompted by the recognition of a new species from the inland deserts. Since
the genus was described in 1829-30 from specimens collected at Shark Bay
in 1818, it has received little attention, being simply enumerated or briefly
described in such works as Bentham, Flora Australiensis (1866) and Lemee,
Dictionnaire descriptif et synonymique des genres de plantes phanerogames
Lamarchea
Lamarchea Gaudichaud-Beaupre in Freycinet, Voy. Gran, et Phys., Bot. 483, tab, 1 10 (text
March 1830, tab. Sept. 1829). Named after M. Lamarche, " capitaine de vaisseau, ex-
lieutenant en pied de fUranie. temoignage d'attachement ”.
Endlicher. Gen. PI. 2:1237 (1836); Bentham & Hooker, Gen. PI. 1:704 (1865); Bentham,
FI. Austral. 3:123 (1866); Bentham, Notes on Myrtaceae, J. Linn. Soc., Bot. 10 (1867);
Lemee. Diet. 3:930 (1931). Lamarchea Reichb. Consp. 175 (1828).
Shrubs with the bark on older stems in papery layers. Young shoots
and flowers with an indumentum of simple hairs. Leaves scattered, flat
or terete, coriaceous. Flowers solitary, scattered, on stems of previous season
or older stems, the buds covered by imbricate deciduous scales. Hypanthium
campanulate with 5 imbricate calyx-lobes. Petals free, longer than the calyx-
lobes. Stamens united in a slightly down-curved tube which is divided into
5 lobes more deeply so on the lower side, the two uppermost lobes the longest,
the lowermost one the shortest, each with about 10-15 marginal hirsute
filaments; anthers versatile. Style filiform, inserted in a depression on the
convex summit of the ovary; stigma small, capitate. Ovai^ sunken in calyx-
tube, 3-celled, with numerous ovules ascending from axile peltate placentae.
Fruit woody, persistent, obovoid, the calyx-lobes, petals and staminal tube
deciduous after flowering. Seeds ± oblong-cuneate, angular, black or brown,
many infertile.
Type Species: Lamarchea hakeifoUa Gaud.
Distribution: Australia: Western Australia and Northern Territory: central
west coast; Victoria, Gibson and southern Great Sandy Deserts (Figure 1).
Bentham (1866) described the ovules as “descending from a peltate
placenta ”, but in fact the placentae are basal. Although closely allied to
the large genus Melaleuca, Lamarchea has a distinctive appearance due to the
solitary flowers with monadelphous stamens. The staminal tube is slightly
curved with unequal lobes, giving a somewhat zygomorphic flower, while the
tube and filaments are hirsute. The flowers have the general aspect of some
species of Calothamnus, which has basifixed anthers. When flowering is
270
over, the petals fall first, followed shortly by the staminal tube and later the
calyx-lobes.
The leaves of Lamarchea sulcata sp. nov. appear to be unique in the Myrta-
ceae in being sulcate. Terete leaves are common, e.g. in Melaleuca and Cah-
thamnus, and they are sometimes singly-channelled on one side, but no other
species with sulcate leaves has been recorded.
Figure I — Distribution of Lamarchea species. O — L- hakeifotia Gaud van hakeifolia
(the localities are close together and cannot be shown individually at this scale). • L. hakei
folia var. brevi folia var. nov. H — L. sulcata sp. nov.
Until recently the genus was considered monotypic, although the former
Government Botanist of Western Australia, the late C. A. Gaidner, recognised
the short-leaved variant of T. hakeifolia as a distinct taxon but did not publish
it These two taxa occur within 50 miles of the coast between the Murchison
River and Shark Bay, so the occurence of another species in the inland deserts
is of interest phytogeographically. A similar situation but in a reverse direction
is in the genus Philotheca (Rutaceae): the recently-described P. tuhiflora A S.
George occurs on the south-western edge of the Great Victoria Desert, while
Ihe other few species are found near the coast in Queensland and New South
Wales A number of other genera are distributed chiefly in wetter parts of
temperate or tropical Australia with a few outlying desert species, e.g. Lomandra
and Thvsanotus (Liliacaeae), Xanthorrhoea (Xanthorrhoeaceae), Kennedia
(Papilionaceae), Hihhertia (Dilleniaceae), Baeckea and Calytrix (Myrtaceae),
Microcorys (Lamiaceae) and Stylidium and Levenhookia (Styhdiaceae).
271
Key to Species
la. L. hakeifolia var. hakeifolia
lb. L. hakeifolia var. brevifolia
2. L. sulcata
I. Leaves flat, 3 (5) — nerved
2. Leaves mostly 3-5 cm long
2. Leaves mostly 1-2 cm long
1 . Leaves terete, sulcate
1. Lamarchea hakeifolia Gaud, in Freycinet, Voy. Uran. et Phys., Bot. 483,
tab. 110 (text March 1830, tab. Sept. 1829). Gaudichaud’s spelling of liakeae-
folia has been altered to hakeifolia in accordance with Art. 73 Recommendation
73 G (c) of the International Code.
Type: in Novae Hollandiae ora occidentali: baie des Chiens-Marins ( - Shark Bay), Gaiidi-
ihund. Iso: FI.
la. var hakeifolia (Figure 2)
A shrub to 5 m, the older stems with bark decorticating in papery layers.
Young shoots densely appressed-sericeous, becoming glabrous, with sub-basal
scale-leaves which are narrow-elliptic, obtuse, up to 7 mm long, pubescent
outside, several-nerved in centre, margins scarious. Leaves oblong to narrow-
lanceolate, sometimes broader in upper half, abruptly narrowed to an acute
apex, shortly (1-2 mm) petiolate, flat, (2-) 3-5 (-6-5) cm long, 4-8 mm wide,
with 3 main nerves, more prominent below, and a marginal or sub-marginal
nerve. Flowers usually on the older stems from which the leaves have fallen,
at first orange- or yellow-green, turning dull red. Buds covered with deciduous
ovate obtuse scales which are ± 2 mm long and pubescent outside. Pedicel
1-2 mm long, pubescent. Hypanthium 3 •5-4' 5 mm long including upper free
part, hirsute: calyx-lobes elliptic-ovate, 4-6 mm long, shortly pubescent with
spreading hairs on both sides, thick but with thin, fimbriate margins. Petals
and staminal tube on a slightly raised narrow disc. Petals oblong-obovate,
obtuse, 11-17 mm long, appressed-pubescent outside in upper half, otherwise
glabrous, thin, the margins almost scarious, ciliate. Stamens 30-43 mm long,
the tube deeply divided, base glabrous, upper part and filaments hirsute except
at apices; anthers linear, 1-1-5 mm long. Style 35-50 mm long, slender,
glabrous, stigma ± 0-75 mm diam. Summit of ovary densely pubescent.
Fruit sessile, depressed-obovoid, 9-12 mm long, 10-12 mm wide, smooth or the
base slightly corky, orifice 3-4 mm diam., truncate or very shortly 5-lobed.
Seed d- oblong — narrow — cuneate, variously angular, 2-3 mm long, shining.
Distribution: Western Australia; central west coast, just south of Shark Bay.
Peron Peninsula, Shark Bay, 26 Aug. 1931, C /«. Gardner 2547; Shark Bay road, 5 miles
N. of Tamala turnoff. 26 Aug. 1969, A. S. George 9559; Shark Bay Road, Oct. 1966, IV. Roger-
son 295; Nilemah Station, Hamelin Pool, 28 Aug. 1931, C. A. Gardner and IV. E. Blackall
538; 15 miles NE of Tamala Station, 13 Oct. I960, 5. Davies (all at PERTH). (Figure 1).
This variety grows in red sand in tall scrub. The main flowering period is
August to October, and new growth begins at this time.
The isotype at FI is in leaf only, with a single detached fruit. Gaudi-
chaud’s illustration shows flowers and buds among the leaves, but it is possible
that this was a “ reconstructed ’’ specimen. The flowers are easily detached in
drying and in this variety they are usually on the old stems only. The illus-
tration also shows the lobes of the staminal tube of equal length with 7-11
filaments each, whereas the upper two are the longest, the lowest one the
shortest, with the other two intermediate, and all have about 10-15 filaments.
lb. var. brevifolia A. S. George, var. nov. (Figure 3)
A varietate typica foliis 1 2 cm longis, 3 -5 mm latis, (loribus plerumque folia ramulorum
juniorum interpositis, calycis lobis 3-5-4-5 mm longis, petalis 10-13 mm longis, staminibus
25-35 mm longis, stylo 25-35 mm longo, fructibus 8-10 mm longis 9-10 mm latis,’ differt.
272
w
Figure 2 Lcur.archea hakeifoUa Gaud. var. Iiakeifolia. Peron Peninsula. Gardner 2547.
Tvne- Near 417 mile peg. North West Coastal Highway ( ± 100 miles N of Geraldton), West-
ern Ausfral it ”1 \m. A. S. George II229. Holo:PERTH, iso : PERTH. AD, BRI,
CANB, K, MEL, NSW, RSA.
Differs from the typical variety in having leaves I -2 cm long, 3-5 mm wide.
Flowers generally among the leaves of the younger branchlets. Calyx lobes
3 . 5 - 4 -5 mm long. Petals 10-13 mm long. Stamens 25-25 mm long. Style
25-35 mm long. Fruit 8-10 mm long, 9-10 mm wide.
Distribution: Western Australia, central west coast between Murchison River
and Shark Bay. up to 50 miles inland.
32 miles
3238 (in fruit)
N of Murchison River. NW Coastal Highway, I8 Feb. I962, A. S. George
; No. 2 Tank, Shark bay (road), I7 Sept. I941, C. A. Gardner 6006 (in fruit);
273
41 1 mile peg, NW Coastal Highway, 3 March 1966, E. M. Scrymgeour 344 (in fruit); 410 mile
peg, NW Coastal Highway, 19 Dec. 1962, F. LuUfitz 1962 (a) (in flower): 35 km NE of Kal-
barri, Murchison River, 18 May 1968, J. Bannister (in fruit) (all at PERTH). (Figure 1).
This taxon differs from the var. hakeifoHa mainly in the shorter leaves,
but the flowers and fruit are also slightly smaller. The indumentum on the
young shoots tends to be more dense and spreading, and persists for a longer
time, sometimes up to a year.
It grows in sandy loam in tall scrub, but occurs to the south of the area
where var. hakeifoHa is found and farther inland. It flowers from November
to January, somewhat later than the typical variety.
One collection at PERTH appears intermediate between the varieties —
7 miles S. of Wannoo, N.W. Coastal Highway, 16 Sept. 1968, M. E. Phillips
Figure 3 — Lamarchea hakeifoHa Gaud. var. brevifoiia var. nov. Holotype — George 11229
274
(duplicate of CBG 025843). It has leaves 2-3 cm long and fruit 8-10 mm long
and 9-10 mm wide. Further collecting in the country between the known areas
of distribution of the two varieties may reveal further intermediates from one
to the other.
2. Lamarchea sulcata A. S. George sp. nov. (Figure 4).
A hakeifolia Gaud, habitu breviori (ad 2 m alt.) patent!, foliis teretibus 5-sulcatis
dilTert. Folia (5-) 15-30 ( 30) mm longa, pungentes. Hypanthium ± 4mm longum, calycis iobi
4-5 mm long!, /’cta/o 10-13 mm longa. 25-40 mm longa. 30-40 cm longus.
FructKs 8-10 mm longus, 7-8 mm latus.
Type: 12 miles E of Todd Range, Gunbarrel Highway, Gibson Desert, Western Australia
(lat. 25‘’43'S, long. 126 II'E), 2 Oct. 1966, A. S. George 8211. In gravelly loam with scattered
Acacia, Eremophiia and Trinclia. Holo: PERTH, iso: PERTH, AD. BRI, CANB, K, MEL,
NSW.
A spreading shrub to 2 m tall with many stems. Bark on old stems in
papery layers. Young shoots closely hirsute, becoming glabrous except for a
few hairs in the sulcae of the leaves. Leaves terete, silicate with 5 grooves,
pungent, (5-) 15-30 (-35) mm long, slightly narrowed into a short petiole.
Flowers scattered on older stems from which leaves have fallen, green and red.
Buds covered by imbricate scales which are narrow to broad-ovate, hirsute
outside, glabrous inside, deciduous. Hypanthium campanulate, densely
hirsute, j; 4 mm long, calyx lobes spreading ovate, obtuse, 4-5 mm long, thick
with somewhat scarious margins, glabrous within. Petals broadly oblong,
concave, obtuse, 10-13 mm long, thin with scarious margins, hirsute outside,
glabrous within. Stamens 25-40 mm long, the tube glabrous outside at the
base, densely hirsute above including the filaments, inside hirsute throughout;
lobes with + 15 marginal filaments; anthers 1 -5 mm long. Style 30-40 mm
long, glabrous; stigma small. Summit of ovary densely hirsute. Fruit al-
most sessile, obovoid, 8-10 mm long, 7-8 mm diam., smooth, dark brown,
orifice ± 5 mm diam., slightly undulate. Fertile seeds few, + oblong, angular,
the outer ones curved on outer surface. 2 mm long, black; sterile seeds narrow,
dark brown.
Distribution: Western Australia and Northern Territory: Victoria, Gibson
and southern Great Sandy Deserts.
W.A. — 9 miles N of No. 12 Well. Cannins? Stock Route, 5 Sept. 1942, H. M. Wilson II
(PERTH); 5 miles N of Camel Well, NE of Wiltma, 8 Sept. 1958, N. H. Speck 1.399 (CANS,
PERTH); 3 miles W. of Yamarna H.S., E. of Eaverton, 2 Julv 1963, A. S. George 4629
(PERTH).
N.T. — South of the Davenport Hills (26 miles E of State border on road from Giles, W.A.
to Sandy Blight Junction), 25 July, 1967 A. S. George 8920 (PERTH, NT. RSA). (Figure 1.)
This species is readily distinguished from L. hakeifolia by its lower, spread-
ing habit and the remarkable silicate leaves. Ft has been collected in several
habitats, viz. on a sandstone ridge (Wilson II), on the crest of a mudstone
escarpment (Speck 1399), on a gentle gravelly slope (George 8211) and on the
sides of sand dunes (George 4629 and 8920). It forms small populations in
widely separated localities. Flowering is apparently dependent on rainfall, as
with most desert plants; Wilson 1 1, collected in September is in flower and bud,
but Speck 1399, collected in the same month of a different year, is in bud only;
George 821 1, (October) is in full flower, as also is George 8920 (July); however,
4629 (July) has fruit only, with no sign of buds.
The Northern Territory record is the first for the genus outside Western
Australia and is an interesting addition to the central Australian flora [G.
Chippendale, Check List of Northern Territory Plants, in Proc. Linn. Soc
N.S. Wales 96, 4: 207-267 (1972)].
276
A taxonomic revision of the genus Tecticornia
(Chenopodiaceae)
By P. G. Wilson
Abstract
An account is given of the genus Tecticornia and of its relationship to other
genera. Two species are described as new; they are T. verrucosa and T. arhorea.
The third species is provided with a new combination as T. austraiasica (based
on Halocnenuiiti auslralasiciim Moq.), The nomenclatural confusion which
has surrounded this name is outlined.
Introduction
The genus Tecticornia is a member of the tribe Salicornieac in the family
Chenopodiaceae. This tribe contains those succulent chenopod genera with
apparently jointed, leafless stems and usually spike-like inflorescences.
The delimitation of genera within this tribe is notoriously diflicult and, as
far as Australian plants are concerned, the available generic names have been
somewhat haphazardly applied by difl'erent authors. From this confusion
the genus Tecticornia does stand apart, for it has, since its inception, been
recognised as being a taxon distinct from the other members of the Australian
Sa/icornieae. This recognition has persisted in spite of the fact that descriptions
of its supposedly diagnostic characters which have been provided by various
authors, are so general as to encompass several genera, or so incorrect as to
render them useless. These errors are nevertheless understandable for the
earlier workers had only dried material for examination, and for an adequate
study of the plants in the flowering stage fresh or pickled material is essential.
Fortunately, through the efforts of several workers mentioned in the ack-
nowledgements, this material has been made available.
Systematic Notes
The genus Tecticornia was placed by Ulbrich (1934) along with Halo-
stachys and Halocnemitm in the subtribe Halostachinae of the tribe Salicornieae.
This systematic arrangement was based on the presence in all three genera of
free opposite and decussate bracts, a character not in this case of phylogenetic
significance, for in habit, and in perianth and seed morphology, Tecticornia
is quite different from the other two.
Free opposite bracts are also found consistently in several Australian
species of Arthrocnemum, and a state of transition from free to united bracts
may be found in other species of that genus. This character cannot therefore
be used by itself to separate the subtribes or genera of the Salicornieae found
in Australia.
The characters which, when taken together, serve to distinguish the genus
Tecticornia from other Australian members of the tribe are: ( I ) the free opposite
and decussate bracts which have fleshy upturned outer margins; (2) the pair
of laterally placed tepals that are eventually free from each other and from the
surrounding bracts; and (3) the disintegration of the spike, when mature, into
its separate components of bracts, tepals, pericarp, and seed. All Tecticornia
species are annuals or short-lived perennials (while in other Australian genera
the species are either long-lived perennials or shrubs), and all have glaucous,
cylindrical articles.
277
Distribution and Habitat
All of the three known species of Tecticornia are native to Australia. Two
are endemic there while the third {T. australasica) is found also in Java and New
Guinea. They are entirely tropical or sub-tropical in distribution (north of
latitude 27°S) except for a single record of T. verrucosa from south of latitude
33°.
Two species, T. australasica and T. verrucosa, grow along the coast in
mudflats above the normal diurnal tidal level. Germination of the seed under
these conditions appears to be dependent upon the seasonal rain leaching the
salt from the upper soil layers. T. arborea has been collected a few miles
from the sea on claypans but it is not found directly on the coast. Both this
species and T. verrucosa also occur far inland where they form monospecific
stands on non-saline claypans, or, in the case of T. verrucosa, also on moderately
saline flats. None of the species occurs around the inland salt-lakes where
other members of the Salicornieae are so common.
Aboriginal Use
The seeds of both T. verrucosa and of T. arborea were at one time used by
Aborigines as food. Further notes on this subject appear under these two
species.
TECTICORNIA Hook. f.
Hook.f. in Benth. et Hook.f., Gen. PI. 3:65 (1880); Black, Trans. Roy. Soc. S. Austral. 43:366
tab. 37 (1919); UIbnch in Engler et Prantl. Nat. Pflanzenfam. ed. 2. I6c:548 (1934)- Backer in
van Steenis, FI. Males, ser. 1. 4:103 (1949).
Annual or short-lived perennial, herbaceous to suffruticose, apparently
leafless plants. Branches apparently Jointed and succulent due to the opposite
leaves enveloping the stem in a cylindrical sheath; leaf-blade minute or absent.
Inflorescence of terminal and lateral spike-like cymes consisting of 3(5) flowers
in the axils of opposite and decussate rows of free (or almost free) bracts.
Bracts closely imbricate, standing vertical to spike axis, broadly deltoid to
semicircular, the outer margin fleshy with an upturned rim. Flowers herma-
phrodite, concealed within bracts and standing vertical to spike axis, free or at
first loosely united to upper bract and to each other; perianth succulent, of
two laterally placed, plano-convex tepals which are free or slightly united along
lower margin towards base, the flat sides pressed together and’ enclosing the
ovary; stamen solitary, abaxial; ovary thin-walled; style slender, weak, with a
pair of filiform stigmas. Fruiting perianth: tepals completely free and, when
dry, areolate and hyaline. Utricle obovoid, with membraneous pericarp.
Seed erect, endospermous; testa thinly coriaceous; embryo on upper (adaxial)
side, slender, curved; radicle inferior; seed dispersed on the drying and sub-
sequent complete disintegration of the spike, perianth, and pericarp.
Type species: T. cinerea (F. Muell.) Baill. [ T. australasica (Moq.) P. G
Wils.].
Origin of name; From the Greek words tectum, a roof, and cornu a horn.
Presumably alluding to the manner in which the outer margin of the bracts
forms a covering to the spike. Probably also to emphasize a relationship with
the genus Salicornia.
1 .
1 .
2 .
2 .
Key to Species
Spikes predominantly terminal; seed ovate with flat sides, shortly aculeate along upper
margin j j australasica
Spikes lateral, or both terminal and lateral; seed neither flat nor aculeate
Spikes lateral, sessile and decussate; seed ± plano-convex, verruculose 2. T. verrucosa
Spikes terminal and lateral (but not sessile); seed bluntly ellipsoidal, smooth
3. T, arborea
278
Figure 1 — Tecticornia verrucosa: A, spike; B, seed; C, T.S. of seed. Tecticornia arborea
D, spike; E, flower from above, F from below, and G from side; FI, seed; I, T.S. of seed
J,’l.S. of seed.
(A-C from Maconochie 1036; D-G from Wilson 8577; H-J from Aplin B20).
279
1. Tecticornia australasica (Moq.) P. G. Wilson comb. nov.
Hahcnemiim aiistralasicum Moq., Chenop. Enum. 110 (1840). — Arthrocnemiim aiisiralasiciini
(Moq.) Moss apud Adamson, j. South Afr. Bot. 20:19 (1954), as to combination only. —
Salicortiia australasica (Moq.) Eichler. Taxon 12:296 (1963), as to combination only. — Type:
■' In nova Hollandia " (holo: P, photo seen, labelled " Nouv. Hollande— Port de Georges ”).
H. ciiiereum F. Muell., Fragm. 1:140 (1859). — Saliconiia cinerea (F. Muell.) F. Muell. ex
Benth., FI. Au.stral. 5:203 (\&10). — Tecticornia cinerea (F. Muell.) Baillon, Hist. PI. 9:185
(1887). — Type: " In locis salinis terrae Arnhem's Land et Australiae centralis " (syn: " Sturt's
Creek. Central Australia”, F. Mueller, MEL 36057 and 36058; " Point Pearce and Sturt’s
Creek ”, F. Mueller. K, n.v., compared by D. .1. McGillivray).
Succulent herbaceous to suffruticose annual, 12-40 cm high. Lower
branches frequently procumbent, upper ones erect; internodes narrowly cylind-
rical, 7-15 mm long, bluish green and glaucous, apex shortly bilobed with a
broad scarious margin. Spikes predominantly terminal (occasionally also
lateral, in which case they are sessile and at right angles to stem), narrowly
ovoid and ca. 1 cm long to narrowly cylindrical and up to 4 cm long, 5-7 mm
diam. Bracts semicircular in shape. Flowers 3-5 in each bract (the lateral
flowers sometimes sterile), ca. 2-5 mm long, at first fused with upper bract
near base, otherwise free. Tcpals laterally appressed, obovoid, acute, plano-
convex in transverse section, at first united below (abaxially) towards base,
otherwise free. Sraininal filament slender, weak; anther oblong-cordate, ca.
I mm long. Seed ovately discoidal, ca. 1-5 mm long; testa dark brown to
black with about 10 indefinite rows of small, grey, translucent and aculeate
tubercles along upper margin, the lower margin sparsely aculeate and the
sides almost smooth. (Figure 2,4.)
Distribution: Coastal tropical Queensland and the Northern Territory (and
Western Australia?). Java and New Guinea. (Figure 7.)
Northern Territory: Fog Bay 12 48' S, 130 21' E. S. T. Blake 16764 (BRI); Coast, 5 mi N of
Finnis R., N. Byrnes 1686 (NT); Port Darwin, Holtze 355 (MEL); 21-8 mi NW of Mount-
norris Bay, G. Chippendale 8168 (NT); East Alligator R., N. Bvrnes 918 (NT); Oenpelli
12 18' S. 133 4' E, R. L. Specht 1 182 (BRI).
Queensland: Karumba, Burke Dist. G. IF. Trapnell 200 (BRI); Cape York, B. Hyland 2501
(BRI); Trinity Bay, W. Hill 5 (MEL); Townsville, N. Michael 474 (BRI): Giru, 19" 31' S,
147 6' E. Nov. 1968, J. Burry (BRI); Caley Valley via Bowen, II Nov. 1959, H. J. Laverv
(BRI).
New Guinea: Terr, of Papua: 8 mi N of Kapa Kapa, R. Schodde 2736 (BRI); 7 mi NW of
Hisiu village, P. ./. Darhyshire 892 (BRI). West Irian: Merauke, H. S. McKee 1703 (NSW).
Ecology : Found on mudflats in coastal or near coastal situations, some-
times in association with mangroves. Mr. N. Byrnes has supplied the
following notes which are derived from his personal observations: “ The plant
is an annual, germinating at the end of the ‘ wet ’. It seldom grows much
larger than 18 in. though some clumps can be dense. It varies between slightly
procumbent to almost prostrate and begins to flower in August. It is normally
found on the salt pans at the back of the mangroves along tidal creeks in pure
communities or associated with Artbrocnennim sp.”.
Apparently it is not able to withstand strongly saline conditions but
grows well in coastal flats seasonally inundated by fresh water.
Notes: The taxonomy of this species has been confused due to the poor material
the earlier workers had available to them and also to the fact that the short
description of Halocnemum aiislralasicuni provided by Moquin was insufflcient
for the name to be used subsequently with any certainty. J. D. Flooker (1880)
equated the name with Saliconiia quinqueflora Bunge ex Ung.-Sternb. but did
not make any formal transfer. This synonymy was also accepted by C. E.
Moss (1954) who considered the species to belong to the genus Arthrocnemum
and made the necessary new combination. Moss typified the species by a
collection made by Robert Brown (" no. 3080 ” at BM), supposedly in the
280
Figure 2 — Tecliconiia ausiralasica: A, spike; B, L.S. of spike; C, flower from below, D from
above, and E from side; F, seed; G, T.S. of seed. (A-E from Byrnes 1950; F-G from J.
Burry, Qld.).
97093 — ( 3 )
281
East Indies, and applied the name to a South African plant which was later
described by H. R. Tolken (1967) as a new species, Arthroaiemiim clecumhetis.
A similar typification was accepted by Hj. Eichler (1963) who transferred
H. australasicum to Saliconiia and placed in synonymy under it S. australis
Solander ex Bentham (1870).
The course followed by Eichler relied on the assumption that the names
//. australasicum Moq. and S. australis Benth. were both based on S'. incHca
[non Willd.] R. Br. (since the last name was cited by Moquin and Bentham in
synonymy under their respective species), and therefore that Bentham's name
was a synonym of Moquin's. The citation provided by Moquin was as follows :
“ Salicornia Imlica R. Brown. Prodr. nov. Holl. 1. p.41l ? non 'Willd. (v.s. in
herb. Mus. Paris) This would appear to state that Moquin considered it
possible that the plant described by R. Brown under the name “ S. indica
Willd.” was in fact the same as his own species, i.e. as H. australasicum. Moquin
had not, however, seen R. Brown's material, nor did he cite it. H. australasi-
cum must therefore be typified by the specimen in Paris which Moquin had
studied and which he had also used to draw up his de.scription.
J. M. Black (1921) attempted to resolve the identity of this type specimen
and published a description of the spike and seed from notes provided for him
by Prof. Lecomte in Paris. Here Black assumed that it was probably related
to Arthrocnemum lylei (Ewart et White) Black, which assumption was accepted
and supported by Tolken (1967). The fact that Black w'as also unable to
apply the name satisfactorily w'as partly due to the still inadequate description
and partly to the confusion of localities. In his original paper of 1840 Moquin
gave as the plant's origin only “ In nova Hollandia". Subsequently (1849) he
added “ ad portum Georgii ", which was interpreted by both Bentham and
Black as being King George Sound, a locality on the south coast of Western
Australia. A photograph of the type of H. australasicum shows that it con-
sists of two portions of stem, one of which has a short lateral branch attached.
Fortunately the actual specimen had previously been studied by Dr. L. A. S.
Johnson who recognised it as being conspecific with Tecticornia cinerea. Mr.
J. Garrick, while working in Europe as the Australian Botanical Liaison Officer,
compared the seed with that obtained from a recent collection of the species
and found them to be the same.
The locality given by Moquin, and which also appears on the herbarium
label as “ Port de Georges ”, is therefore either an error (as the species is
tropical in distribution), or the name refers to a place other than King George
Sound.
The only collection recorded from an inland situation was that made by
F. von Mueller in 1856 along Sturt Creek, which is in the north-east of Western
Australia. I have seen one other collection of a Tecticornia from this area ;
this was made by Dr. J. S. Beard in 1968 at Lake Doman (approximately 70
km west of the southern end of Sturt Creek) and is of T. verrucosa. Although
T. verrucosa has been confused with T. australasica, all of Mueller's specimens
belong in fact to the latter species. These collections constitute therefore the
only inland record for T. australasica (if the locality data are correct), and also
the only record for Western Australia.
The application of the name Salicornia australis Sol. ex Benth. (1870)
depends on its lectotypification. Solander applied the name (in manuscript)
to a specimen collected by J. G. and J. R. Forster in New Zealand. This
specimen was also referred to by Bentham and, if made the lectotype, the
name would probably become a synonym of S. quinqueflora Ung.-Sternb.
(1866).
The combination " Tecticornia cinerca " has been variously attributed to
J. D. Hooker (1880), F. M. Bailey (1901), and J. M. Black (1919). Hooker
never in fact published the combination and the first person to do so was
282
283
Figure 3—Tecticornia verrucosa, x approx. 1/4 (A. L. Payne s.n.). Figure 4 — 7i?cr/cor;;/u a;Hrra/ax/ea, x approx. 3/4 (N. Byrnes 1950).
apparently H. E. Baillon in 1887. It was subsequently made by B. Daydon
Jackson in Index Kewensis 2:1041 (1895) and then independently by both
F. M. Bailey and J. M. Black.
2. Tecticornia verrucosa P. G. Wilson sp. nov.
Spicae oppositae et decussatae, ad axem rami verticales, sessiles, breviter cylindraceae,
10-20 mm longae, 6 mm latae. Semina late et obtuse elliptica vel suborbicularia, patelli-
formia (leviter concavo-convexa), ca. 1-8 mm longa, verruculosa.
Type: 9 mi SE of Rabbit Flat 20’ 15' S, 130° E, Northern Territory, small erect shrub with
several stems to 8 in. high, 26 May 1970, ./. R. Maconochie 1036 (holo: PERTH, iso: K,
MEL, NSW).
Annual or short-lived perennial, to 40 cm high, branching at the sometimes
woody base. Lower branches often decumbent and giving rise to erect stems.
Articles cylindrical (to obovoid), 10-14 mm long, 5-6 (18) mm diam., pale
bluish green (or pale red), glaucous; apex truncate with minute lobes; margin a
narrow scarious rim. Inflorescence of opposite and decussate lateral spikes
along upper portion of branches (occasionally a very small terminal spike is
present), each opposite pair being separated from neighbouring pairs by a
single internode. Spikes sessile, standing at right angles to branch axis,
shortly cylindrical, 10-20 mm long, 6 mm diam.; bracts at first loosely adherent
laterally to each other along basal margins, eventually free. Flowers in triads,
free from each other and from lower bract, united in basal half to upper bract,
the lateral florets loosely united in basal half to lateral bracts. Perianth obovoid,
obtuse, ca. 2 mm long, elliptical in transverse section; tepals free below (abaxi-
ally) but lightly united adaxially in lower half, eventually completely free.
Staminal filament eventually subulate; anther oblong-cordate, ca. 1-3 mm long
(sometimes aborted). Style slender, the stigmas filamentous and ca. 3-5 mm
long. Seed broadly and bluntly elliptical to suborbicular, plano-convex to
slightly concavo-convex, ca. 1-8 mm long, verruculose around upper margin,
grading to ± smooth below, brown to black, surrounded by and adherent to
the transparent membranous pericarp. (Figure lA-C, 3).
Distribution: Tropical and sub-tropical North-West and Central Australia.
Also known from a single locality in the southern portion of Western Australia
near Lake Grace. (Figure 7).
iVestern Australia: Derby, R. D. Royce 3336 (PERTH); ibid., A. 5. George 6532 (PERTH);
13 mi ESE of Wyndham township, R. A. Perry 2559 (NSW, NT); 13 mi WSW of Yeeda
Station, Kimberleys, M. Lazarides 6586 (PERTH); Noblys Well, E. Kimberley, Aug. 1906,
iV. K Fitzgerald (NSW 126370); Lake Doman, J. S. Beard 5601 (PERTH); Dovers Hills,
ca. 30 mi S of Lake Mackay, 13 June 1962, J. Long (NT, PERTH); SE of Lake Anec, Lat.
24 S, June 1958, W. H. Moyle (PERTH); Boolathanna Station N of Carnarvon, T. E. H.
Aplin B14 (PERTH); 4 mi W of Dovers Hills, northern Gibson Desert, A. S. George 9003
(PERTH); ca. 13 km ESE of Mt. William Lambert, Sept., 1971. D. Lowry (PERTH); Lake
Bryde, ca. 20 mi WSW of Newdegate, T, E. M. Aplin 4793 (PERTH).
Northern Territory: 1 mi SE of Rabbit Flat, 130“ E, 20“ 15' S, J. R. Maconochie 1031 (PERTH);
75 mi W of The Granites, 21 Mar. 1963, A. J. Mahood (NT 10257); 3 mi NE of Lake Mackay,
G. Chippendale 3391 (BRI, MEL, NSW, NT, PERTH).
Ecology: Found on coastal mudflats and also inland on fresh water claypans
and slightly saline flats.
Aboriginal Use: A note attached to a specimen collected by J. Long from about
30 mi S. of Lake Mackay in 1962 states that the plant was used by natives as a
food. A further account of its use as a food occurs in a “ Report on health
and nutrition of natives from Rawlinson Range to Lake MacDonald, 1958 ”
compiled by J. J. Elphinstone for the Department of Public Health, Western
Australia. In this paper Mr. W. H. Moyle contributed the following comment :
“ Mungiba {Tecticornia cinerea)
284
285
This plant usually grows in salt pan country. The only patch we saw was
in a fresh water clay pan S.E. of Lake Anec and almost on the 24° latitude.
The plant is not known to natives in the Warburton-Rawlinson area, but it is a
favourite source of food for those further north. The fleshy bulbs containing
small seeds are eaten direct from the plant and have a pleasant, slightly salty
flavour.”
Specimens of the plant referred to have been lodged with the Western
Australian Herbarium and are of T. verrucosa.
Notes: There is a slight variation in the shape and degree of verrucosity of the
seed. In the northern collections the seed is narrower and bears more numerous
but smaller verrucosities compared with seed from plants collected in inland
localities.
The coastal form behaves as an annual. Its seed germinates after im-
mersion in fresh water following the summer rains. In inland localities the
plant has sometimes the appearance of being two or more years old.
3. Tecticornia arborea P. G. Wilson sp. nov.
Spicac lerminales et laterales, pedunculatae (baud sessiles), cylindraceae vel leviter fusi-
formes, 10*20 mm longae, 7 mm latae. Semina late ellipsoidea, obtusa ca. I -5 mm longa,
laevia.
Type: Bulli Bulli Clay Pan, Glen Station, ca. 60 km NW of Cue, Western Australia; erect
pine-like plants up to 172 cm high (average about 135 cm), well-spaced on hard reddish brown
clay, the only species present; 5 Aug. 1969, P. G. Wilson 8577 (holo: PERTH, iso; AD, B,
BRI, CANB, GH, K, L, MEL. NSW).
Figure 7 - Distribution of Tecticornia species. Tecticornia anstraiasica • T. arborea -|-
T. verrticosa Q.
286
Erect pyramidal-shaped plant, loosely branched and with a single stem,
to ca. 150 cm high, woody towards base. Articles on branchlets cylindrical.
truncate. 1-3 cm long, 5-12 mm diam., green to purple, glaucous. Spikes
terminal to main and lateral branches (not sessile), cylindrical or slightly I'usi-
form, 10-20 mm long, ca. 7 mm diam. smooth, bracts free, reniform. Flowers
protogynous and arranged in triads, free from each other and from bracts.
Perianth deltoid-shaped from above (broadest at ape.x), ca. 3 mm long, dorsi-
ventrally flattened at apex, distal half thick and succulent, proximal portion
thin; tepals separated on upper (adaxial) side, united in basal two-thirds on
lower side, eventually free, slightly imbricate at apex. Stamina! filament
subulate; anther cordate-oblong, ca. 2 mm long. Seed broadly ellipsoidal,
rounded at both ends. ca. I -5 mm long, with long axis vertical to spike axis;
testa smooth, reddish brown, glossy; embryo near upper margin of seed sur-
rounded by copious endosperm. Pericarp, eventually tearing at base and
forming a loose cap to the ripe seed. (Figure I D-J, 5, 6).
Distribution: Known from only two localities in subtropical Western Australia
between approximately the 24° and 27° of latitude. (Figure 7).
W end of Weld Range, 15 .lune 1961, .S'. Davies (PERTH); ibid., Apr. 1969, Pat. Lindsey
(PERTH); 23 mi E of Kalli, N. H. Speck 1066 (PERTH); Boolathanna Station N of Carnarvon,
T. F. H. Apiin. B. 20 (PERTH); Carnarvon, Apr. 1962 ./. N. Hnichinson (PERTH).
It is probable, in spite of the varied locality data, that the plant has in
fact been found in only two places, the one about 60 km north-west of Cue and
the other north of Carnarvon.
Ecology: In the two localities where it has been recorded it grows in fresh-water
clay pans as a monospecific stand.
Notes: This species is remarkable because of its erect tree-like habit. It appears
to be capable of living for several years although precise data on this point are
lacking. A vigorous stand was observed at the type locality in August 1969
but a year later all the mature plants were dead and no young ones were found.
Presumably regeneration from seed would have commenced with the first
good rain of the new season.
Aboriginal Use: The seed is large compared with that of other samphire species,
and was at one time collected by aborigines for use as a food. They gave it
the name of “ bulli bulli ”.
Acknowledgements
Several people have substantially assisted in the study of the genus Teeti-
cornia. Mr. N. Byrnes of the Northern Territory Administration at Darwin
has sent fresh material of T. australasica, while Mr. J. R. Maconochie of the
Northern Territory Administration, at Alice Springs and Mr. A. L. Payne
of the W.A. Department of Agriculture at Derby have both provided fresh
rnaterial of T. verrucosa.
During his period as Australian Botanical Liaison Olficer at Kew Mr.
D. J. McGillivray arranged for a photograph of the type of Halocnemum
australasicum (which is in Paris) to be sent to me and also compared the syn-
types at Kew of Halocnemum cinereum with recent collections of this species.
Mr. J. Garrick, who later occupied the same position, kindly compared the
seed on the type specimen of H. australasicum with that from several species
in the Salicornieae.
287
References
Bailey, F. M. (1901). Queensland Flora 1261.
Black, J. M. (1919). A revision of the Australian Salicornieae. Trans. Rov. Soc. S. Austral.
43:355-367.
(1921) Additions to the flora of South Australia No. 19. Trans. Roy. Soc. S.
Austral. 45:8-10.
FIooker, J. D. (1880). Halocnemum and Tecticornia In Bentham and Flooker, Genera
Plantarum 3:65.
Moquin-Tandon, a. (1849). Flalocnemuni in A.P. de Candolle, Prodromus Systematis
Naturalis 13 (2): 149-150.
Tolken, H. R. (1967). The species of Arthrocnemum and Salicornia in Southern Africa-
Bothalia 9;255-.307.
288
Taxonomic notes on Western Australian species of Pityrodia,
Beaufortia and Verticordia
By A. S. George
Abstract
The correct application of the name Pityrodia axillaris (Endl.) Druce is
shown and a new combination. P. terminaiis (bndl.) comb. nov. is provided for
the species to which the former name was incorrectly applied. Beaufortia
macrostemou Lindl. var. ineana Benth. is raised to specific rank.
The following names are shown to be correct for three species of Verticordia
which have commonly been otherwise referred to -
V. hrachypoda Turcz, V. chrysostat hys Meisn. and V. hehusH S. Moore.
Introduction
These notes are intended to provide the correct names for several com-
monly-collected plants. Preliminary work is being done towards a revision
of the genus Verticordia but it is unlikely that this or similar studies of Beau-
fortia and Pityrodia will appear for some years.
Pityrodia R. Br. (Dicrastylidaceae)
Pityrodia terminaiis (Endl.) A. S. George comb. nov.
Basionym: Dasvmalia terminaiis Endl., in Endl. et Fenzl. Nov. Stirp. Dec. 12
(15 May 1839).'
Type: Interior of S.W. Australia, Roe. Holo.- W.
Synonyms.' Pityrodia racemosa (Turez.) Benth., FI. Austral. 5:50 (1870) —
Qitoya? racemosa Turez., Bull. Soc. Imp. Nat. Mosc. 2:194 (1863). Types:
Western Australia, Drummond CoW. 3:141 and Coll. 5:73. Isosyntypes of 5:73
at PERTH. K.
Chloanthes stachyodes F. Muell., Fragm. Phyt. Austral. 5:50 (Julyl865)—
Qiiova stachyodes F. Muell. l.c. nomen pro syn. Type: In fissuris rupium
graniticarum Australiae occidentalis interioris longitudine 113° 45', latitudine
30° 15'. .^coll.
Chloanthes grandifiora Moldenke, Phytologia 2:310 (22 April 1947).
Type: Western Australia, J. Mauritzon, Sept. 1936. Holo.' S.
This is the species which for many years has been incorrectly referred to as
Pityrodia axillaris (Endl.) Druce, and before the publication of that combination
as P. racemosa (Turez.) Benth. Examination of the types and original descrip-
tions shows that P. axillaris is the species which was recently described as P.
spectahilis C. A. Gardn. (see below'). The other names cited above all apply
to the same species, and Endlicher's epithet is the earliest available.
Pityrodia axillaris (Endl.) Druce, Rep. Bot. Exch. Cl. Brit. Isles 1916:640
(\9\l)-~Dasymalia axillaris Endl., Nov. Stirp. Dec. 11 (15 May 1839). Type:
Interior of S.W. Australia, Roe. Holo: W.
Synonym: Pityrodia spectahilis C. A. Gardn., Journ. Roy. Soc. W. Austral.
47:63 (1964). Type: in distr. Irwin prope Buntine in arenosis lutosis, fl., m.
Decern., Gardner 12023. Holo: PERTH.
As mentioned above, the name Pityrodia axillaris has been incorrectly
applied to the species which should be called P. terminaiis. This led C. A.
Gardner to describe as a new species the plant to which the name was originally
applied. Endlicher de.scribed the stamens as included, but this was due to
his examining unopened buds, the dissections of which are still present on the
Holotype sheet.
v7093-(4)
289
Beaufortia R. Br. (Myrtaceae)
Beaufort ia iiicana (Beiith.) A. S. George comb, et stat. nov.
Basionym: Beaufortia macrostemon Lindl. var. incana Benth., FI. Austral.
3:167 (1866). Type: Swan River (Western Australia), Drummond. Holo." K.
This differs in several important respects from typical Beaufortia macros-
temon Lindl., and deserves specific rank. It is a tall rigidly-branched shrub of
1-2 m without a lignotuber, i.e. killed by fire and regenerating from seed. The
leaves are narrow-linear, 1 -nerved or nerveless, and densely appressed-pubes-
cent, broader on new growth. The calyx-lobes are triangular, obtuse, fz 1 mm
long( about half the length of the petals), densely pubescent, with inconspicuous
nerves. The fruiting capsule is h 2 mm diam. Beaufortia macrostemon is a
shrub of 20-50 cm with many relatively slender branches arising from a lig-
notuber and regenerating from this after fire. The leaves are narrow-linear to
narrow-lanceolate, 1-nerved and often with 1-2 lateral nerves, the latter some-
times marginal, and the indumentum of short and long hairs, very spreading
and much less dense than in B. incana. The calyx-lobes are almost subulate,
T 3 mm long (as long as the petals), sparsely hirsute-pubescent, and 3-nerved.
The fruiting capsule is ^ 3 mm diam.
Beaufortia incana is frequent on gravelly heaths between Pingelly and
Nyabing, while B. macrostemon occurs in open woodlands along the Darling
Scarp between Mogumber and Pinjarra.
Verticordia DC. (Myrtaceae)
Attention is drawn to the correct names for the following species of Verti-
cordia:
Verticordia brachypoda Turcz., Bull. Soc. Nat. Mosc. 20, 1 :158 (1847). Types:
(Western Australia) Drummond Coll. 3:28 (BM, FI, K); Gilbert 30 (n.v.).
Synonyms: Verticordia fimbripetala Turcz., Bull. Soc. Nat. Mosc. 22, 1:19
(1849). Type: Drummond 4:47 (FI, K, NSW, W.).
Verticordia stylotricha Diels, Bot. Jahrb. 35:403 (1904). Type: pr. Tammin
in arenosis fruticulosis apertis flor, m. Oct., Diels 5052 (BM, E, K, P.).
Note that V. multiflora Turcz. is a distinct species differing from V. brachy-
poda in the bright yellow flowers and in floral morphology. The two were
considered conspecific by W. E. Blackall in How to Know Western Australian
Wildflowers Vol. 1, p. 278 (1954), but the varietal combination cited there
(V. multiflora var. stylotricha) has never been validly published.
Verticordia chrysostachys Meisn., Journ. Linn. Soc. Bot. 1:41 (1857). Type:
(Western Australia) Drummond CoW. 6:46 (Holo: NY, iso: BM, CGE, E, FI, K,
P. W).
Note that this is the correct spelling, not chrysostachya as used by Bentham
and subsequent workers.
Verticordia helmsii S. Moore, Journ. Linn. Soc. Bot. 34:190 (1898). Type: Nr.
Gnarlbine, fl. Nov., S. Moore (K, NY); ad Warangering, Helms “in Herb.
Kew ” (n.v.).
Synonym: Verticordia adenocalyx Diels, Bot. Jahrb. 35:404 (1904). Type: pr.
Karalee inter fruticeta praecipue Acaciarum in arenosis, fl. m. Nov., Diels
5566 (n.v.).
Although the type of V. adenocalyx has not been found, the description,
and area and time of collection, agree well with those of V. helmsii.
Publication date of Volume 1 Number 2
The publication date of Volume I Number 2 was 5 May, 1971.
97093/4;72-lM
WILLIAM C. BROWN, Government Printer, Western Australia
CONTENTS
Page
A revision of the Australian species of Hybanthus Jacquin (Violaceae).
By Eleanor M. Bennett 218
Four new taxa of Eucalyptus from Western Australia.
By M. I. H. Brooker 242
Studies in the genus Acacia — 1.
By B. R. Maslin 254
New species and subspecies of Casuarina in Western Australia.
By L. A. S. Johnson 261
New taxa and new combinations in Western Australian Pittosporaceae.
By Eleanor M. Bennett 266
A revision of the genus Lamarchea Gaudichaud (Myrtaceae: Leptosnerm-
oideae).
By A. S. George 270
A taxonomic revision of the genus Tecticornia (Chenopodiaceae).
By P. G. Wilson 277
Taxonomic notes on Western Australian species of Pityrodia, Beaufortia
and Verticordia
By A. S. George 289
Publication date of Volume 1 Number 2 290
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